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A Monograph of the Lichen Genus
Pseudoparmelia Lynge (Parmeliaceae)
Z/ :■!
MASON E. HALE, JR.
SMITHSONIAN CONTRIBUTIONS TO BOTANY
NUMBER 31
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SMITHSONIAN CONTRIBUTIONS TO BOTANY • NUMBER 31
A Monograph of the Lichen Genus
Pseudoparmelia Lynge (Parmeliaceae)
Mason E. Hale, Jr.
ISSUE©
SEP 3 1315
SMITHSONIAN INSTITUTION PRESS
City of Washington
1976
ABSTRACT
Hale, Mason E., Jr. A Monograph of the Lichen Genus Pseudoparmelia Lynge
(Parmeliaceae). Smithsonian Contributions to Botany , number 31, 62 pages, 18
figures, 1976. — A world monograph of the genus Pseudoparmelia is presented.
There are 76 species, most occurring in dry subtemperate to tropical areas and
especially concentrated in the arid scrub lands of South Africa, Australia, and
Brazil. The genus is characterized by an unusually high number of species with
divaricatic acid and related orcinol depsides. The following new species are
described: P. callichroa Kurokawa, P. concomitans Hale, P. conlabrosa Hale,
P. dahlii Hale, P. neoquintaria Hale, P. subambigua Hale, P. subamplexa Hale,
and P. venezolana Hale. The following new combinations are made: P. cor-
rugativa (Kurokawa and Filson) Hale, P. exornata (Zahlbruckner) Hale, P.
gerlachei (Zahlbruckner) Hale, P. papillosa (Lynge ex Gyelnik) Hale, P.
rodriguesiana (Hue) Hale, P. schistacea (Kurokawa and Filson) Hale, P. scoto-
phylla (Kurokawa) Hale, and P. spodochroa (Kurokawa and Filson) Hale.
Official publication date is handstamped in a limited number of initial copies and is recorded
in the Institution’s annual report. Smithsonian Year. Series cover design: Leaf clearing from the
Katsura tree Cercidiphyllum japonicum Siebold and Zuccarini.
Library of Congress Cataloging in Publication Data
Hale, Mason E.
A monograph of the lichen genus Pseudoparmelia Lynge
(Parmeliaceae)
(Smithsonian contributions to botany ; no. 31)
Bibliography: p.
Includes index
Supt. of Docs, no.: SI 1.29:31
1. Pseudoparmelia. I. Title. II. Series: Smithsonian Institution. Smithsonian contributions
to botany ; no. 31.
QKI.S2747 no. 31 [QK585.P2] 581'.08s [589'.1] 75-619375
Contents
Page
Introduction 1
Morphology 2
Chemistry 5
Ecology and Habitats 6
Formation of Morphs and Chemical Populations 7
Phytogeography 8
Taxonomic Treatment 10
Doubtful and Rejected Names 56
Literature Cited 58
Index 61
iii
A Monograph of the Lichen Genus
Pseudo par melia Lynge (Parmeliaceae)
Mason E. Hale Jr.
Introduction
This world monograph of the lichen genus
Pseudoparmelia is part of my continuing effort to
revise the parmelioid genera. Although these genera
are usually lumped under the collective name Par-
melia, recent studies with the ‘scanning-electron
microscope have demonstrated that species previ-
ously classified under Parmelia fall into two distinct
groups, one with a paraplectenchymatous upper
cortex and very frequently with pseudocyphellae,
typified by Parmelia saxatilis (L.) Acharius and in-
cluding the P. borreri group, and another with a
palisade plectenchymatous upper cortex and an
overlying pored epicortex but never with pseudo-
cyphellae (Hale, 1973a).
The epicorticate group includes several hetero-
geneous elements differing, for example, in marginal
ornamentation (cilia, bulbate cilia), rhizine branch-
ing patterns (simple or dichotomously branched),
and lobe configuration and width (broad apically
rotund to narrow obtuse lobes). These elements are,
in my opinion, biologically isolated and represent
good genera, as follows: Bulbothrix Hale with bulb-
ate cilia and no cortical pigments (Hale, 1974),
Hypotrachyna (Vainio) Hale with narrow eciliate
lobes and dichotomously branched rhizines (Hale,
1975a), Parmelina Hale with narrow ciliate lobes
(Hale, 1974), Parmotrema Massalongo with broad
rotund lobes (Hale, 1965a), Pseudoparmelia Lynge
with narrow eciliate lobes, Relicina (Hale and Ku-
rokawa) Hale with bulbate cilia and usnic acid in
Mason E. Hale, Jr., Department of Botany, National Mu-
seum of Natural History, Smithsonian Institution, Washing-
ton, D.C. 20560
the cortex (Hale, 1975b), and Xanthoparmelia
(Vainio) Hale with narrow eciliate lobes and usnic
acid in the cortex (Hale, 1974).
As with Hypotrachyna (Hale, 1975a) and Relicina
(Hale, 1975b), much of the initial work on the
chemistry of Pseudoparmelia was completed before
thin-layer chromatography came into use in 1965.
Thus it has been necessary to reinvestigate the
chemistry of many specimens. Even with the applica-
tion of the most recent techniques, however, this
monograph can be considered definitive only in the
broadest sense. Fresh lichen collections are now be-
ing made every year in exotic regions, especially in
Africa, Australia, and South America, by botanists
and more significantly by professional lichenologists,
who had not previously collected outside temperate
areas and who choose to identify the specimens
themselves rather than send them to a monographer.
Under these circumstances, any monograph will be
incomplete in two respects. For one, information on
new records not available for study will be omitted,
records that could profoundly affect interpretations
of phytogeography. The older specimens now pre-
served in herbaria are a pitifully inadequate repre-
sentation of lichen floras. Secondly, unpublished
new species which are sure to be found in a genus as
large as Pseudoparmelia obviously cannot be in-
cluded in the keys, and this lack makes species
identification more difficult for lichen students. This
monograph is offered, then, with full realization of
its limitations but with the hope that it will pro-
vide a base for later workers.
Acknowledgments. — I am much indebted to
curators who sent specimens on loan and often
allowed considerable extensions of loan periods.
1
2
SMITHSONIAN CONTRIBUTIONS TO BOTANY
These include Mr. Peter James (BM), Dr. A. J.
Kostermans (BO), Dr. A. Robyns (BR), Mr. M. S.
Christiansen (C), Dr. W. A. Weber (COLO), Dr. W.
L. Culberson (DUKE), Dr. L. Williams (F), Dr. I.
M. Lamb (FH), Dr. C. E. Bonner (G), Mr. C. E.
Palmar (GLAM), Dr. T. Ahti (H), Dr. H. Schindler
(KR), Dr. J. Poelt (M), Dr. H. A. Imshaug (MSC),
Dr. H. Osorio (MVM), Dr. C. T. Rogerson (NY), Dr.
Hildur Krog (O), Dr. E. F. Warburg (OXF), Mme.
Jovet-Ast (PC), Dr. C. E. Smith (PH), Dr. S. Ahlner
(S), Dr. S. Kurokawa (TNS), Dr. R. Alava (TUR),
Dr. I. Tavares (UC), Dr. R. Santesson (UPS), Dr. H.
Riedl (W), Dr. J. W. Thomson (WIS), and Dr. G.
Cufodontis (WU). Dr. Ove Almborn (LD) lent his
own extensive collections in addition to materials
from BOL, L, PRE, and TRH which he had on
hand for study. Dr. Z. CernohorskJ kindly arranged
for loans from BP at a time when travel in Hungary
was not permitted. The late Dr. H. des Abbayes
(REN) was extremely generous in sending loans as
well as duplicates of type-collections of the new
species he was describing. For allowing me to ex-
amine collections in their private herbaria, I am
especially indebted to Dr. D. D. Awasthi, Dr. Gun-
nar Degelius, Dr. M. Mahu, Dr. T. D. V. Swinscow,
and Dr. Rolf Santesson.
The specimens were photographed by the Smith-
sonian Office of Printing and Photographic Ser-
vices. Scanning-electron microscope photographs
were prepared by the Smithsonian Scanning-Elec-
tron Microscope Laboratory.
Dr. S. Kurokawa assisted with species identifica-
tions, chemical tests, and descriptions prior to 1961,
and his help is gratefully acknowledged. Field
studies have been supported at various times by the
National Science Foundation, National Geographic
Society, Smithsonian Research Foundation, and the
Morden-Smithsonian Fund. Cooperative field
studies were conducted in India with the Maharash-
tra Association for the Cultivation of Science, work-
ing with Dr. P. G. Patwardhan, and in Venezuela
with Dr. M. Lopez-Figueiras of the Universidad de
los Andes. Finally, I wish to thank Dr. Ove Alm-
born for reading the entire manuscript and check-
ing the bibliography and spelling of place names.
Morphology
Thallus. — The thallus is typically foliose, orbic-
ular, 3-15 cm in diameter, and uniformly adnate.
even at the margin. The degree of adnation varies
from nearly subcrustose in P. xanthomelaena (Fig-
ure 18e) to loosely adnate in P. caperata. Compa-
rable eciliate species of Parmotrema have wider,
more or less marginally erect lobes. Lobe configura-
tion is variable. While some species have sublinear,
apically obtuse lobes 0. 5-2.0 mm wide (Figures 6b, d,
lOe, 12 d), an even larger number have subirregular,
apically rotund lobes 2-6 mm wide (Figures 6a, c,
9a, 10c, 12c). Lobes in both types are often contigu-
ous and crowded. There is no marginal ornamenta-
tion in Pseudoparmelia; the margins are smooth and
sometimes black rimmed.
The lower surface of the thallus is either black
or pale brown, and this color difference, although
variable in some of the saxicolous species, can be
an important taxonomic character. Broader lobed
species with a black lower surface, such as the well-
known P. caperata, usually have a narrow brown
zone at the lobe tips. While most species have a
shiny paraplectenchymatous lower surface (Figure
lc), species in the P. amplexa group (P. amplexa, P.
pachydactyla, and P. subamplexa) have a minutely
grained, velvety appearing black lower surface to
the margin (Figure 1 a,b). This is caused by an ir-
regularly thickened, minutely papillose cell layer
sloughing from the lower cortex, a phenomenon
which I have not seen in any other parmelioid
genera.
Rhizines are uniformly simple and produced in
moderate abundance (Figure Id). The only excep-
tion to this rule is P. intertexta, which usually has
pale branched rhizines, not unlike those of some
species of Relicina but not as agglutinated (Hale,
1975b). Broad-lobed species usually lack rhizines
along the margins at lobe tips or at most have a
zone of short papillae. The rhizines are often blunt
with pale, penicillate tips (Reznik et al., 1968).
The internal structure of Pseudoparmelia pre-
sents no unusual features (Figure 2 a,c,f). The upper
cortex has a basic palisade structure (Figure 2d) and
is overlain by a thin pored epicortex (Figure 1 e,f),
as I had previously discovered by using the
scanning-electron microscope (Hale, 1973a). The
medullary hyphae are often conspicuously encrusted
with lichen substances. The lower cortex is para-
plectenchymatous (Figure 2e). In these respects it
is identical with the other epicorticate genera in the
family.
Vegetative Propagules. — The isidia in Pseudo-
NUMBER 31
3
Figure 1. — Morphology of Pseudoparmelia: a, lower surface of P. amplexa (Degelius s. n.)
(X 160); b, enlargement of surface in a (X 1600) with arrow pointing to a papilla; c, lower
surface of P. rutidota ( Degelius A-37) (xl6Q); d, cross section of P. malaccensis to show simple
rhizines ( Hale 30353) (xl40); e, surface of P. rahengensis showing pored epicortex ( Kurokawa
1602) (X 400); f, enlargement of surface in e showing pores and aggregations of crystals of usnic
acid (X 1600). (All photographs taken with the scanning-electron microscope.)
4
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Figure 2. — Morphology of Pseudoparmelia: a, longitudinal cross section of P. venezolana with
arrow in upper right pointing to the epicortex (Hale 45526a) (X 390); b, enlargement of epi-
cortex in a (upper arrow) and upper cells of the cortex (lower arrow) (X 4000); c, longitudinal
cross section of P. spodochroa ( Weber 47220) (X 370); d, enlargement of upper cortex in c
(X 800); e, longitudinal cross section of P. sphaerospora ( Culberson 10324) (x 240); f, enlarge-
ment of lower cortex in e (X 800); g, isidia of P. dahlii (isotype in US) (X 160).
NUMBER 31
5
parmelia are quite uniform. Twenty species (P.
adspersa, P. amazonica, P. annexa, P. arcana, P.
basutoensis, P. caroliniana, P. cinerascens, P. con-
crescens, P. conlabrosa, P. cyphellata, P. dahlii (Fig-
ure 2 g), P. ecaperata, P. ischnoides, P. malaccensis,
P. martinicana, P. neoquintaria, P. salacinifera, P.
scotophylla, P. subtortula, and P. venezolana) have
normal, cylindrical, simple to sparsely branched
isidia. Two additional species (P. pachydactyla and
P. papillosa) have abnormally thickened but other-
wise normal isidia (Figure 14 d). No species have
lobulate, procumbent, or ciliate isidia.
Pustules have been described and illustrated for
Hypotrachyna (Hale, 1975a), and they are more or
less the same in Pseudoparmelia (Figure 7 b), very
thickened, fragile, hollow isidioid structures which
usually break open apically. Some then produce
soredia, while in other cases no soredia form. Seven
species (P. baltimorensis, P. eruptens, P. geesterani,
P. owariensis, P. pustulescens, P. schistacea, and P.
zimbabwensis) have largely nonsorediate pustules.
One species (P. raunkiaeri) has heavily sorediate
pustules.
Soralia occur in 14 species: P. alabamensis, P.
aptata, P. caperata, P. carneopruinata, P. crozalsi-
ana, P. cryptochlorophaea, P. epileuca, P. gerlachei,
P. labrosa, P. leucoxantha, P. soredians, P. subam-
bigua, P. subamplexa, and P. texana. They are usu-
ally orbicular and discrete except in P. caperata,
which has coalescent, more diffuse soralia. All are
strictly laminal except for P. leucoxantha (Figure
136), where the soralia are largely submarginal. It is
perhaps noteworthy that all of the sorediate species,
excepting P. alabamensis, are corticolous; sorediate,
obligately saxicolous species are very rare in the
genus.
The remaining 32 species lack any vegetative
propagules. The older lobes, however, may become
lobulate or heavily rugose, although, as pointed out
above, no species have true laminal lobulae (phyl-
lidia) or lobulate isidia. Virtually all of these species
are fertile.
Apothecia. — Apothecia, found so far in 47 of
the 76 species, are extremely uniform. They are
sessile to subpedicellate and 1-3 mm (rarely to 6
mm) in diameter. The disc is always imperforate.
The asci contain eight spores, and the bulk of the
species have spores in the range of 6-10 ^m wide
and 7-13 pm long. Pseudoparmelia caperata and
P. concomitans have the largest spores, up to 24 p.m
long.
Chemistry
The chemical constitutents of Pseudoparmelia
were first summarized by Hale and Kurokawa (1964)
on the basis of microcrystal tests. I have retested
most of the species with thin-layer chromatography,
using solvent systems A and B of C. Culberson
(1972) and in large part confirming the earlier re-
sults. The lichen substances in the genus are listed
below under the presently accepted classification
with number of species containing each substance
in parentheses.
Aliphatic Acids
Caperatic acid (5)
Protolichesterinic acid (3)
Others (2)
Aromatic Compounds
Orcinol Series
para-Depsides
Divaricatic acid (9)
Evernic acid (1)
Gyrophoric acid (4)
Lecanoric acid (4)
Obtusatic acid (1)
Perlatolic acid (2)
meta-Depsides
Crytochlorophaeic acid (1)
Sekikaic acid (1)
Depsidones
Neoloxodic acid (2)
Norlobaridone (3)
yj-Orcinol Series
para-Depsides
Atranorin (55)
Barbatic acid (1)
4-0-Demethylbarbatic acid (1)
Depsidones
Constictic acid (5)
Fumarprotocetraric acid (1)
Norstictic acid (2)
Physodalic acid (2)
Protocetraric acid (25)
Salazinic acid (7)
Stictic acid (8)
Succinprotocetraric acid (1)
Dibenzofuranes
Usnic acid (20)
6
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Anthraquinones
Skyrin (2)
Undetermined (3)
Unidentified substances presumed to be depsi-
dones are known in P. chalybeizans (“chalybeizans”
unknown), P. chapadensis, P. exornata and P. papil-
losa (“conformata” unknown), and P. neoquintaria
(“quintaria” unknowns). Unidentified depsides or
orcinol depsidones are probably present in P. van-
derbylii. Unidentified yellow pigments have been
reported for P. arcana, P. baltimorensis, P. condyl-
oides, P. corrugativa, P. cyphellata, P. rutidota, and
P. sphaerospora, but none of these are accompanied
by triterpenoids. Details on these will be found in
“Taxonomic Treatment.”
Divaricatic acid, perlatolic acid, and sekikaic acid
are all accompanied by 1 to 3 weaker, as yet uniden-
tified spots falling below the main acid. Perlatolic
acid may occur mixed with (or even be replaced by)
stenosporic acid, but these two substances are not
resolved in the solvent systems employed here.
Gyrophoric acid always occurs as an accessory with
protocetraric acid, never alone, and norstictic is ac-
companied by salazinic acid or stictic acid.
While I hope to discuss the chemical features of
Pseudoparmelia in relation to the other parmelioid
genera in a separate article, several chemical traits
stand out. The abundance of orcinol series depsides,
especially divaricatic acid, is noteworthy and in fact
characterizes the genus. The absence of the com-
moner orcinol series depsidones such as alectoronic
acid is significant, as is also the absence of echino-
carpic acid, galbinic acid, lichexanthone, and triter-
penoids, and the rarity of barbatic acid and
norstictic acid.
Ecology and Habitats
Knowledge of the ecology of any lichen group can
enhance our understanding of difficult species com-
plexes, and it is best if this knowledge is gained
through firsthand experience in the field. Herbar-
ium labels often omit important data on general
vegetation, substratum, and elevation. While I have
collected specimens of Pseudoparmelia in various
parts of the world, I have never visited Africa or
Australia, two very important centers for the genus.
It is still possible, however, to draw a general pic-
ture of the ecological requirements of the genus.
It is evident, for example, that Pseudoparmelia is
best developed in temperate, subtropical, and arid
tropical regions at low to mid elevations. This is in
contrast to the strong boreal requirements of Par-
melia, and the highland tropical preferences of
Hypotrachyna (Hale, 1975a). Only two species, P.
baltimorensis and P. caperata, grow as far north as
the southern edge of the boreal forest, although in
the Southern Hemisphere P. gerlachei occurs in
subantarctic localities. At the same time only four
species occur with any frequency in humid lowland
tropical forests: P. sphaerospora in the Americas
and Africa and P. dahlii, P. intertexta, and P. mal-
accensis in Southeast Asia.
Pseudoparmelia occurs most abundantly in the
arid desert or sclerophyll scrublands of Africa and
Australia. In these two regions alone nearly 30
obligately saxicolous species are known and most
of them are endemic. The approximately 40 ob-
ligately corticolous species in the genus occur
mostly in open deciduous secondary forest or in
disturbed sites such as pastures, parks, and botanical
gardens. About 8 species occur on either rocks or
trees, the most typical being P. texana. These are
generally habitats of great environmental stress with
intense insolation and prolonged drought.
Pseudoparmelia competes with two other par-
melioid genera: Parmotrema, which is mainly corti-
colous, and Xanthoparmelia , a saxicolous group. In
South Africa Parmelia subgenus Melanoparmelia
is also well devolped on rocks, and as a result Pseu-
doparmelia, Xanthoparmelia, and the Melanopar-
meliae often have similar lobe configuration and
close thallus adnation. They all have simple rhi-
zines, lack marginal cilia, and rarely produce sore-
dia. It may represent an example of convergent
evolution for three genera in these stressed environ-
ments.
Another possible example of. convergent evolu-
tion is represented by the three usnic-acid contain-
ing species in the humid lowland forests, P. dahlii,
P. intertexta, and P. malaccensis. They occupy the
same habitats (in particular canopy branches in
dipterocarp forests) and have the same narrow, ap-
pressed, sublinear lobes and yellow pigmentation
(usnic acid) as Relicina (Hale, 1975b) but lack the
marginal bulbate cilia.
NUMBER 31
7
Formation of Morphs and Chemical Populations
A detailed discussion of morph formation and
speciation in Hypotrachyna (Hale, 1975a), which
applies equally well to Pseudoparmelia, has already
been presented. Briefly, one assumes that nonfertile
isidiate and sorediate morphs have evolved from
fertile nonisidiate or nonsorediate parent species
and that lacking any means of sexual reproduction
the morphs tend to conserve chemical traits. A par-
ent morph may be traced rather easily, may already
be modified through the usual genetic changes, or
may have become extinct and now be represented
only by the vegetative morph. While application
of the morph concept may be somewhat arbitrary in
actual practice, it has offered a reasonable explana-
tion of species evolution in several foliose genera.
There are seven theoretically possible combina-
tions of parents and vegetative morphs if we con-
sider only soredia and isidia (Figure 3). Pustules,
which in any event are not common, would be
considered as an offshoot of isidia. In Pseudopar-
melia the largest category is the 16 fertile species
which, as far as we can determine from herbarium
collections now available, have not produced any
vegetative morphs. Some will undoubtedly be found
eventually, but the presumption is that these spe-
cies (P. callichroa, P. caribaea, P. chapadensis, P.
concomitans, P. condyloides , P. corrugativa, P.
hypomilta, P. inornata, P. lecanoracea, P. rodrigue-
siana, P. rupicola, P. rutidota, P. somaliensis, P. sub-
tiliacea , P. vanderbylii, and P. violacea), most rather
rare, have not yet evolved successful vegetative
morphs.
Referring again to Figure 3, the next largest
category is 12 isidiate morphs for which no corre-
sponding parents have been discovered. These in-
clude P. adspersa, P. amazonica, P. arcana , P. caro-
liniana, P. cinerascens, P. dahlii, P. geesterani, P.
neoquintaria, P. owariensis, P. rahengensis, P. salac-
inifera , and P. venezolana. We might assume in this
instance that the morphs evolved from a nonisidiate
parent which has not been able to survive and pro-
pagate or which has not yet been discovered.
Coexisting parents and isidiate morphs (but with
no sorediate morph) are represented by the follow-
ing nine species pairs: P. exornata - P. papillosa,
P. intertexta - P. malaccensis (tentative), P. molyb-
diza - P. annexa, P. prolata - P. basutoensis, P.
Parent (16)
(lacking) (lacking)
Parent (9)
Isidiate (9) (lacking)
Parent (4)
(lacking)
Isidiate (3) Sorediate (3)
(lacking)
Isidiate (12) (lacking)
(lacking)
(lacking) Sorediate (4) (lacking)
Parent (2)
Sorediate (6)
Isidiate (2) Sorediate (2)
Figure 3. — Patterns of morph formation in Pseudoparmelia
with number of species in each category in parentheses.
spodochroa - P. scotophylla, P. tortula - P. subtor-
tula, P. xanthomelaena - P. ischnoides, and P.
zambiensis - P. ecaperata.
Soredia are comparatively rare in Pseudoparme-
lia. There are four parent species with correspond-
ing sorediate morphs: P. epileuca - P. schelpei, P.
ferax - P. gerlachei, P. inhaminensis - P. crozalsiana
(tentative), and P. scrobicularis - P. carneopruinata.
Six sorediate species, P. alabamensis, P. aptata, P.
cryptochlorophaea, P. leucoxantha, P. soredians,
and P. subambigua, appear to have no nonsorediate
parent.
Two groups of species have the full complement
of morphs: P. amplexa - P. pachydactyla - P. sub-
amplexa and P. nairobiensis - P. concrescens - P.
texana (including as well P. eruptens). A third
series, P. rutidota - P. baltimorensis (coarsely pustu-
late)-!3. caperata (sorediate), is another possibility
but the species here all diverge rather widely from
each other in some details.
A final group of four species, P. martinicana - P.
raunkiaeri and P. conlabrosa - P. labrosa, represent
isidiate-sorediate morph pairs but have no fertile
parent extant.
Of the remaining species in the genus, P. geester-
ani, P. pustulescens , P. schistacea, and P. zim-
8
SMITHSONIAN CONTRIBUTIONS TO BOTANY
babwensis are pustulate and have no obvious
relatives.
There is a particularly interesting group of saxi-
colous species in South Africa, including P. molyb-
diza (lecanoric acid), P. spodochroa (salazinic acid),
P. tortula (norlobaridone), and P. vanderbylii (un-
known depsides). They are so similar externally
that chemical tests are needed to separate them.
While this group may have evolved from a common
parent, the chemical evolution has followed very
different pathways. The presumptive isidiate
morphs of three of these (P. annexa, P. scotophylla,
and P. subtortula) are also sufficiently similar that
chemical tests are required for positive identifica-
tion.
If one subscribes to the concept of vegetative
morphs, it then follows that morphologically identi-
cal but chemically different vegetative morphs have
evolved from different parents through parallel evo-
lution and are therefore polyphyletic (W. Culberson,
1973). Pseudoparmelia texana (divaricatic acid) and
P. aptata (perlatolic acid), for example, differ in
distribution, P. texana being pan-temperate whereas
P. aptata occurs only in the Old World. The two
acids involved are para-depsides, divaricatic acid
having a C3H7 side chain and perlatolic acid C5Hn
side chains (Figure 4). Another chemical pair, P.
owariensis (divaricatic acid) and P. pustulescens
(sekikaic acid), is confined to the Old World and
CHoCH
-CO— O-,
-OH
-OH
-COOH
R
ch3o
c3h7
OH
CO— 0-,^S,-C00H
OH CH3ol J-C3H7
Figure 4. — Comparison of molecular structures of acids in
Pseudoparmelia: Divaricatic acid (R = C3H7) and perlatolic
acid (R = C6Hu) (above) and sekikaic acid (below).
appears to be sympatric. The acids are particularly
interesting since sekikaic acid is a meta-depside
and divaricatic acid the homologous para-depside
(Figure 4).
Rarer species with chemical differences used as
taxonomic characters include P. dahlii (lecanoric
acid) and P. rahengensis (barbatic acid group) in
Southeast Asia. Pseudoparmelia rutidota, a common
species in Australia and southwestern United States
into tropical America, contains protocetraric acid;
a virtually indistinguishable population, P. exor-
nata, contains protocetraric acid with the “confor-
mata” unknown and occurs principally in Uruguay.
Other close relatives include P. ferax, which con-
tains physodalic acid, and P. callichroa, which con-
tains a fatty acid.
The presence or absence of usnic acid is the main
criterion for recognizing P. nairobiensis (usnic acid
absent in the cortex) and P. zambiensis (usnic acid
present) and their corresponding isidiate morphs,
P. concrescens and P. ecaperata.
Phytogeography
The Pseudoparmelia floras of various geopolitical
units are listed below. Many countries are so poorly
collected, of course, that no meaningful floristic
comparisons can be made. However, the United
States, West Indies, Venezuela, South Africa, and
perhaps Brazil and Australia are all rather well
known. Several lichenologists are now working in
East Africa and Australia, and their studies, when
completed, will fill in many gaps.
North America
Canada: P. baltimorensis, P. caperata.
United States: P. alabamensis, P. amazonica, P. baltimoren-
sis, P. caperata, P. caroliniana, P. crozalsiana, P. crypto-
chlorophaea, P. martinicana, P. rutidota, P. salacinifera, P.
sphaerospora, P. texana.
Mexico and Central America
Mexico: P. amazonica, P. carneopruinata, P. caroliniana, P.
crozalsiana, P. leucoxantha, P. martinicana, P. raunkiaeri,
P. rutidota, P. salacinifera, P. sphaerospora, P. texana.
Guatemala: P. caperata, P. sphaerospora.
Honduras: P. amazonica, P. carneopruinata, P. caroliniana,
P. cryptochlorophaea, P. salacinifera, P. texana.
Nicaragua: P. caroliniana.
Costa Rica: P. carneopruinata, P. caroliniana, P. sphaerospora,
P. texana.
Panama: P. caroliniana, P. sphaerospora, P. texana.
NUMBER 31
9
West Indies
Bahamas: P. sphaerospora.
Cuba: P. amazonica, P. caribaea, P. caroliniana, P. salacini-
fera, P. sphaerospora.
Jamaica: P. carneopruinata, P. caroliniana, P. cryptochloro-
phaea, P. raunkiaeri.
Hispaniola: P. caperata, P. caroliniana, P. cryptochlorophaea,
P. inornata, P. martinicana, P. raunkiaeri, P. sphaerospora,
P. texana.
Puerto Rico: P. amazonica, P. martinicana.
Lesser Antilles and Virgin Islands: P. caribaea, P. crypto-
chlorophaea, P. inornata, P. martinicana, P. raunkiaeri, P.
sphaerospora.
Trinidad: P. amazonica.
South America
Colombia: P. amazonica, P. carneopruinata, P. salacinifera.
Venezuela: P. amazonica, P. caperata, P. carnepruinata, P.
P. caroliniana, P. cryptochlorophaea, P. martinicana, P.
salacinifera, P. scrobicularis, P. sphaerospora, P. texana,
P. venezolana.
Ecuador: P. caroliniana.
Peru: P. caperata, P. rutidota, P. sphaerospora, P. texana.
Bolivia: P. rutidota.
French Guyana: P. caribaea, P. sphaerospora.
Brazil: P. amazonica, P. caperata, P. carneopruinata, P. caro-
liniana, P. chapadensis, P. cinerascens, P. crozalsiana, P.
cyphellata, P. exornata, P. hypomilta, P. leucoxantha, P.
papillosa, P. rupicola, P. rutidota, P. salacinifera, P. scrobi-
cularis, P. sphaerospora, P. texana.
Uruguay: P. carneopruinata, P. crozalsiana, P. exornata, P.
papillosa, P. rutidota, P. texana.
Paraguay: P. cinerascens, P. scrobicularis.
Argentina: P. caperata, P. carneopruinata, P. crozalsiana, P.
gerlachei, P. papillosa, P. rutidota, P. scrobicularis, P. sore-
dians.
Chile: P. caperata, p. ferax, P. gerlachei, P. labrosa, P. rutid-
ota, P. soredians, P. subambigua, P. texana.
Europe and Africa
Europe: P. caperata, P. caroliniana, P. carneopruinata, P.
crozalsiana, P. soredians.
Tunisia: P. caperata.
Guinea: P. ecaperata, P. pustulescens, P. rodriguesiana, P.
sphaerospora.
Ivory Coast: P. caroliniana, P. ecaperata, P. malaccensis, P.
owariensis, P. rodriguesiana, P. texana.
Cameroon: P. sphaerospora.
Ethiopia: P. aptata.
Somalia: P. somaliensis.
Uganda: P. annexa, P. arcana, P. caroliniana, P. concrescens,
P. molybdiza, P. nairobiensis, P. owariensis, P. pustulescens,
P. somaliensis, P. subtortula, P. texana.
Urundi: P. somaliensis.
Kenya: P. caperata, P. epileuca, P. nairobiensis, P. pachydac-
tyla, P. soredians, P. subtortula, P. texana.
Zaire: P. concrescens, P. crozalsiana, P. sphaerospora.
Rhodesia: P. subamplexa, P. tortula.
Angola: P. amplexa, P. annexa, P. arcana, P. caroliniana, P.
concrescens, P. ecaperata, P. inhaminensis, P. nairobiensis,
P. pustulescens, P. sphaerospora, P. texana.
Malawi: P. ecaperata.
Zambia: P. somaliensis, P. zambiensis.
Tanzania: P. aptata, P. molybdiza, P. nairobiensis, P. so-
maliensis, P. sphaerospora, P. texana.
Mozambique: P. concrescens, P. epileuca, P. eruptens, P.
malaccensis, P. schelpei.
Union of South Africa: P. amplexa, P. annexa, P. aptata,
P. arcana, P. basutoensis, P. concrescens, P. condyloides, P.
crozalsiana, P. eruptens, P. geesterani, P. ischnoides, P.
lecanoracea, P. molybdiza, P. prolata, P. pustulescens,
P. rodriguesiana, P. soredians, P. spodochroa, P. texana,
P. tortula, P. vanderbylii, P. violacea, P. xanthomelaena.
Madagascar: P. rodriguesiana, P. somaliensis, P. sphaerospora,
P. texana.
Asia
India: P. aptata, P. caperata, P. crozalsiana, P. ecaperata, P.
malaccensis, P. pustulescens, P. texana.
Sri Lanka: P. dahlii, P. texana.
Thailand: P. adspersa, P. ecaperata, P. owariensis, P. rahen-
gensis, P. salacinifera, P. texana.
Laos: P. caroliniana
China: P. caperata.
Japan: P. aptata, P. caperata, P. owariensis, P. texana.
Hong Kong: P. owariensis.
Taiwan: P. amazonica.
Philippines: P. adspersa, P. intertexta, P. malaccensis.
Malaysia: P. intertexta, P. malaccensis.
Indonesia: P. aptata, P. intertexta, P. malaccensis, P. texana.
New Guinea: P. intertexta.
Australia and New Zealand
Australia: P. aptata, P. caperata, P. conlabrosa, P. corrugativa,
P. ferax, P. intertexta, P. labrosa, P. neoquintaria, P. ruti-
dota, P. schistacea, P. scotophylla, P. soredians, P. spodo-
chroa, P. subtiliacea, P. texana, P. xanthomelaena.
New Zealand: P. caperata, P. labrosa, P. rutidota, P. soredians,
P. subtiliacea.
Pacific Recion
New Caledonia: P. concomitans.
Hawaii: P. caperata, P. texana.
The greatest concentration of species is found in
South Africa. This region with adjacent countries
has a flora of 35 species, nearly half of the species
in the genus (Figure 5). At least 15 of these are
endemic and most saxicolous. Australia, a region
of similar climate and vegetation, has 16 species, of
which 5 (P. aptata, P. soredians, P. spodochroa, P.
texana, and P. xanthomelaena ) also occur in Africa.
In the New World, Brazil has the most species,
18, of which 6 are endemic. North America has 12
species but only 2 (P. alabamensis and P. balti-
morensis) are endemic there.
Temperate and tropical eastern Asia (India to
10
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Japan) has a rather small Pseudoparmelia flora of
only 14 species, 4 of them endemic and the re-
mainder pantemperate or also widespread in Africa.
In general the Old World and New World floras
are quite distinct. The Old World, however, is
much richer with about 55 species in contrast to
the 30 species so far collected in North and South
America.
Some idea of the abundance of the various spe-
cies can be gained by tabulating the number of
countries (as listed above) where each species has
been reported. This crude method of comparison
does not, of course, take into consideration the
relative sizes of the countries or the intensity of
collecting, but it is unlikely that the order of
abundance will be greatly changed by future col-
lecting. The following species occur in five or more
countries: P. texana (24), P. caperata (17), P. sphae-
rospora (16), P. caroliniana (13), P. amazonica (9),
P. carneopniinata (9), P. crozalsiana (9), P. rutidota
(9), P. salacinifera (8), P. ecaperala (7), P. aptala (7),
P. concrescens (6), P. cryptochlorophaea (6), P.
malaccensis (6), P. martinicana (6), P. soredians (6),
P. intertexta, (5), and P. owariensis (5). Within its
range P. caperata is undoubtedly the the most com-
monly collected foliose lichen, followed by P. caro-
liniana and in Asia P. ecaperata.
Twenty-nine species, nearly half the genus total,
have been found so far in only one country, as can
be determined from the lists of specimens examined
in the species treatment, and while many of these
will be found in other countries with more intensive
collecting, this great a number reflects the high de-
gree of endemism characteristic of the genus.
Taxonomic Treatment
The 76 species of Pseudoparmelia are arranged
below in alphabetic order. All specimens collected
by Hale are deposited in the Smithsonian Institu-
tion (US), and the herbarium acronym is not cited
for these.
The following key is divided into three sections:
isidiate species, sorediate species, and nonisidiate,
nonsorediate species.
Keys to the Species of Pseudoparmelia
P. malaccensis
P. dahlii
P. raliengensis
ISIDIATE
1. Thallus greenish yellow (usnic acid present).
2. Thallus black below with at most a narrow brown zone at the lobe tips.
3. Isidia fine, no more than 0.2 mm thick; medulla P — P. eeaperata
3. Isidia thick and inflated, 0.5-1 mm thick; medulla P+ red.
4. Lower surface velvety black to the margin P. pacliydactyla
I. Lower surface shiny black with a narrow brown zone at the lobe tips.
5. Isidia dense, cylindrical P. papillosa
5. Isidia irregularly scattered, often breaking open apically. . . P. baltimorensis
2. Thallus uniformly pale brown hplow.
6. Medulla P+ red (protocetraric acid).
6. Medulla P-.
7. Medulla C+ red (lccanoric acid)
7. Medulla C- or C+ orange (barbatic acid)
1. Thallus whitish to greenish mineral gray (usnic acid lacking).
8. Collected on trees.
9. Lower surface uniformly pale brown.
10. Isidia moderate to dense, 1 mm high; medulla K+ red (salazinic acid)
P. salacinifera
10. Isidia sparse, 2-3 mm high; medulla K— or K+ yellow (stictic acid)
P. cyphellata
9. Lower surface black toward the center and black at the margin or with a narrow
brown zone at the lobe tips.
II. Isidia inflated, pustulate, 2-3 mm high P. eruptens
11. Isidia not inflated or pustulate, to 1 mm high.
12. Upper surface finely reticulately cracked and strongly white-reticulate
P. caroliniana
12. Upper surface continuous, not white-reticulate.
13. Medulla P-.
14. Medulla C+ red (lecanoric acid) P. conlabrosa
14. Medulla C— (divaricatic acid) P. concrescens
13. Medulla P+ red or orange.
15. Medulla K+ (salazinic acid) P. cinerascens
15. Medulla K-.
16. Fumarprotocetraric acid present P, adspersa
16. Protocetraric acid present.
17. Isidia fragile, crumbling; lobe tips white-reticulate
P. martinicana
17. Isidia normal; lobes not white-reticulate. P. amazonica
8. Collected on rocks.
18. Isidia coarse, pustulate, usually erupting apically.
19. Medulla K+ red (salazinic acid) P. geesterani
19. Medulla K-.
20. Medulla P+ red (protocetraric acid) P. zimbabwensis
20. Medulla P-.
21. Divaricatic acid present P. oivariensis
21. Divaricatic acid absent.
22. Sekikaic acid present P. pustulescens
22. Caperatic acid present P. schistacea
18. Isidia normal, thin and cylindrical, not pustulate.
23. Medulla P+ red or orange.
24. Lobes 2-3 mm wide, apically subrotund P. scotophylla
24. Lobes about 1 mm wide.
25. Medulla K — ; collected in South America P. venezolana
25. Medulla K+ yellow (stictic acid); collected in Africa. P. ischnoides
23. Medulla P-.
26. Lobes 2-4 mm wide, apically subrotund.
12
SMITHSONIAN CONTRIBUTIONS TO BOTANY
27. Upper surface finely reticulately cracked P. caroliniana
27. Upper surface continuous and smooth P. subtortula
26. Lobes 1-2 mm wide, apically obtuse.
28. Lower surface black P. annexa
28. Lower surface uniformly brown or tan.
29. Thallus very closely appressed. P. arcana
29. Thallus adnate to loosely adnate.
30. “Quintaria” unknowns present; collected in Australia
P. neoquintaria
30. “Quintaria” unknowns absent; collected in Africa.
31. Norlobaridone present. P. subtortula
31. Norlobaridone absent. P. basutoensis
Sorediate Species
1. Thallus greenish yellow (usnic acid present).
2. Lower surface uniformly velvety black below to the margin P. subamplexa
2. Lower surface shiny, black at the center and dark brown in a narrow zone at the lobe tips.
3. Medulla K+ yellow turning red (salazinic acid) P. soredians
3. Medulla K-.
4. Medulla C+ red (lecanoric acid), P — . P. subambigua
4. Medulla C — , P+ red or orange.
5. Soralia mostly laminal, becoming diffuse P. caperata
5. Soralia mostly apical or marginal.
6. Soralia generally orbicular; collected in Antarctic regions or the high Andes.
P. gerlachei
6. Soralia elongate along lobe margins; collected in mid-elevation tropical re-
gions P. leucoxantha
1. Thallus whitish to greenish mineral gray (usnic acid absent).
7. Medulla P + orange or red.
8. Upper surface broadly reticulately ridged and foveolate (without lens); stictic acid
present.
9. Lobes subirregular, 2-5 mm wide P. crozalsiana
9. Lobes sublinear, 1-2.5 mm wide P. carneopruinata
8. Upper surface smooth and plane; protocetraric acid present.
10. Soredia coarse, densely produced over the upper surface in coalescing soralia
P. raunkiaeri
10. Soredia farinose, produced in discrete orbicular soralia.
11. Collected on rocks P. alabamensis
11. Collected on trees P. epileuca
7. Medulla P-.
12. Medulla C+ deep red (lecanoric acid). P. labrosa
12. Medulla C— or C+ fleeting pink.
13. Soralia strongly capitate along lobe margins. P. cryptochloropliaea
13. Soralia not capitate, entirely laminal.
14. Divaricatic acid present P. texana
14. Perlatolic acid present P. apt at a
Nonisidiate, Nonsorediate Species
1. Thallus greenish yellow (usnic acid present; P. chapadensis, P. hypomilta, and P. sphaerospora
appear to be greenish yellow but lack usnic acid).
2. Lower surface brown or tan throughout,
3. Collected on trees P. intertexta
3. Collected on rocks P. chapadensis
2. Lower surface black.
4. Lower surface velvety black to the margin P. amplexa
4. Lower surface shiny black and often with a narrow brown zone at the lobe tips.
5. Medulla P-.
6. Collected in Africa P. zambiensis
6. Collected in South America P. callichroa
5. Medulla P+ red.
7. Physodalic acid present P. ferax
7. Physodalic acid absent.
8. Protocetraric acid present P. rutidota
8. Protocetraric acid and the “conformata” unknown present P. exornata
1. Thallus whitish to greenish mineral gray (usnic acid absent; P. chapadensis, P. hypomilta,
P. sphaerospora, and sometimes P. violacea appear to be yellowish green but lack usnic acid).
9. Medulla pigmented purple to yellow.
10. Medulla completely deep purple P. violacea
10. Medulla yellow or orange.
11. Medulla completely pale orange-yellow ( P . condyloides appears yellow but con-
tains no pigment.) P. sphaerospora
11. Medulla pigmented only in the lower half.
12. Collected on rocks P. lecanoracea
12. Collected on trees.
13. Lower surface brown; collected in South America P. hypomilta
13. Lower surface black; collected in Australia P. corrugativa
9. Medulla white.
14. Collected on trees.
15. Lower surface uniformly brown.
16. Medulla P+ red (protocetraric acid) P. somaliensis
16. Medulla P— or P+ faint orange (stictic acid) P. sphaerospora
15. Lower surface black (often with a narrow brown zone at the lobe tips).
17. Medulla P-.
18. Collected in Africa; divaricatic acid present P. nairobiensis
18. Collected in Australia and New Zealand; caperatic acid present
P. subtiliacea
17. Medulla P+ red or orange.
19. Medulla K+ yellow (stictic acid); surface of lobes reticulately ridged
and foveolate (without lens).
20. Lobes 1-2 mm wide; collected in South America. P. scrobicularis
20. Lobes 2-4 mm wide; collected in Africa P. inhaminensis
19. Medulla K— (protocetraric acid).
21. Lobes 1-2 mm wide P. schelpei
21. Lobes 3-6 mm wide.
22. Medulla C— ; collected in the West Indies P. inornata
22. Medulla C+ rose; collected in New Caledonia
P. concomitans
14. Collected on rocks.
23. Lower surface uniformly brown.
24. Medulla P+ orange (salazinic acid).
25. “Chalybeizans” unknown present P. condyloides
25. “Chalybeizans” unknown absent P. spodochroa
24. Medulla P-.
26. Medulla C+ red (lecanoric acid) P. molybdiza
26. Medulla C-.
27. Thallus rather loosely attached on rocks and soil; lobes subelongate.
P. prolata
27. Thallus tightly adnate on rocks; lobes shorter, apically rotund.
28. Norlobaridone present P. tortula
28. Unknown substances present P. vanderbylii
23. Lower surface black (brown only in a narrow zone at the lobe tips).
29. Medulla P+ orange or red.
30. Lobes less than 1 mm wide P. xanthomelaena
30. Lobes 2-4 mm wide.
31. Medulla K— ; collected in the Caribbean region P. caribaea
31. Medulla K+ yellow or red (salazinic acid); collected in Australia
and Africa P. spodochroa
29. Medulla P-.
32. Collected in South America.
P. rupicola
14
SMITHSONIAN CONTRIBUTIONS TO BOTANY
32. Collected in Africa.
33. Medulla C+ red (lecanoric acid) P. molybdiza
33. Medulla C— (divaricatic acid) P. rodriguesiana
Pseudoparmelia
Pseudoparmelia Lynge, 1914:15.
Type-Species. — Pseudoparmelia cyphellata Lynge,
1914:15.
Lynge erected this genus on the basis of a single
collection from Brazil, citing the presence of “pseu-
docyphellae” on the lower surface as the main
difference from Parmeiia. Santesson (1942) demon-
strated conclusively that these are in reality artifacts
caused by rhizines that had been torn off, revealing
the medulla in irregular patches. He did not reject
the genus, as implied in the title of his article, but
regarded it as a synonym of Parmeiia. I do not be-
lieve that one could invoke Article 71 of the Code
of Botanical Nomenclature here to reject Pseudo-
parmelia. The gashes of dislodged rhizines on the
lower cortex are not monstrosities in the accepted
sense. Lynge simply described a new genus on a
mistaken interpretation of the morphology. In any
event. Article 71 was deleted from the Code at the
sessions of the International Botanical Congress
in Leningrad in 1975. The only lichenologist to use
this genus name in practice was Gyelnik (1933),
whose species (Pseudoparmelia aradensis and P.
pseudofallax) have been placed in Cetrelia (Culber-
son and Culberson, 1968).
Pseudoparmelia adspersa
Figure 6a
Pseudoparmelia adspersa (Vainio) Hale, 1974:189.
Parmeiia adspersa Vainio, 1907:168 [type collection: Lem
Ngob, Siam, Schmidt XV (TUR, lectotype; C, isolectotype)].
Parmeiia filipina Hale, 1972a:99' [type collection: Tagaytay,
Cavite, Philippine, Hale 26809 (US, holotype; TNS, UPS,
isotypes)].
Description. — Thallus adnate on trunks, whitish
ashy gray, 5-8 cm broad; lobes subrotund, 3-6 mm
wide, the upper surface plane, continuous or finely
cracked with age, densely isidiate, the isidia mostly
simple, cylindrical, 0.1-0.15 mm high, 0.05 mm in
diameter; lower surface black except for a narrow
dark brown zone along the margins, sparsely to
moderately rhizinate. Apothecia (from Hale 26976)
rare, substipitate, 2-4 mm in diameter, the amphi-
thecium isidiate; spores 8, 5 X 10-12 ^m.
Chemistry. — Cortex K+ yellow, medulla K — ,
C — , KC — , P+ red; atranorin, fumarprotocetraric
acid, and succinprotocetraric acid.
Distribution. — Philippines, Thailand.
Habitat. — Planted Casuarina trees at low to mid
elevation (200-2000 m).
Remarks. — The type collections ( Schmidt XV
and XXXIV) are fragmentary. When I finally re-
solved the chemistry with TLC, I soon discovered
that the species is the same as Parmeiia filipina. Des
Abbayes (1958) compared P. adspersa with Parmeiia
inactiva (Zahlbruckner) Vainio, a much larger am-
phigymnioid lichen containing protocetraric acid.
I had compared it with Pseudoparmelia carolinana
(Nylander) Hale, which contains perlatolic acid
and has a more consistently and conspicuously
cracked cortex.
Specimens Examined. — Philippines: Quezon Province, Hale
26976; Mountain Province, Santos 88 (PHU, US).
Pseudoparmelia alabamensis
Figure 6 b
Pseudoparmelia alabamensis (Hale and McCullough) Hale,
1974:189.
Parmeiia alabamensis Hale and McCullough, 1968:44 [type
collection: St. Clair County, Alabama, Hale and McCul-
lough 24072 (US; isotypes in DUKE, UPS)].
Description. — Thallus closely adnate on rock,
ashy white, 2-4 cm broad; lobes narrow and sub-
linear, contiguous, 0.8-1. 2 mm wide; upper surface
transversely cracked with age, sorediate, the soralia
laminal, capitate; lower surface black, moderately
rhizinate, the rhizines black. Apothecia unknown.
Chemistry. — Cortex K+ yellow, medulla K — ,
C — , KC— or rose, P+ red; atranorin and proto-
cetraric acid.
Distribution. — United States (Tennessee and
Alabama).
Habitat. — On large sandstone rocks in open de-
ciduous forests at about 300 m elevation.
Remarks. — This species is unique among the
North American Pseudoparmeliae. An African spe-
NUMBER 31
15
Figure 6. — Species of Pseudoparmelia: a, P. adspersa ( Hale 26809, holotype of Parmelia filipina
Hale); b, P. alabamensis (McCullough 1842 in US); c, P. amazonica ( Hale 19389); d, P. amplexa
( Gossweiler 3256e in US); e, P. annexa (Almborn 5683, holotype in LD); f, P. aptata (Almborn
6548 in LD). (Scale in mm.)
16
SMITHSONIAN CONTRIBUTIONS TO BOTANY
cies, P. epileuca (Hale) Hale, also has soredia and
protocetraric acid, but it is corticolous and has
broader lobes.
Specimens Examined. — United States: Tennessee, Hale
36927.
Pseudoparmelia amazonica
Figure 6c
Pseudoparmelia amazonica (Nylander) Hale, 1974:189.
Parmelia amazonica Nylander, 1885:611 [type collection:
Santarem, Brazil, Spruce 111 (H, Nylander herbarium num-
ber 35111, lectotype; BM, G, NY, W, PC, isolectotypes)].
Description. — Thallus closely adnate on bark,
5-10 cm broad, light mineral gray and turning buff
in the herbarium; lobes subirregular, apically ro-
tund, 2-6 mm wide, contiguous; upper surface
plane, continuous, moderately isidiate, the isidia
rarely branched, to 2 mm high; lower surface black,
densely rhizinate except for a narrow naked or
papillate zone along the margins. Apothecia rare,
1-3 mm in diameter; spores 8, 8-10 X 13-16 /xm.
Chemistry. — Cortex K+ yellow, medulla K — ,
C — , KC+ rose, P-f red; atranorin and protocetraric
acid.
Distribution. — United States (Florida), Central
America, West Indies, Colombia, Venezuela, Brazil,
Guinea, and Taiwan.
Habitat. — On trees (conifers, palm trees, decid-
uous trees), Opuntia, and on rocks in open forests
at 100-1500 m elevation.
Remarks. — Superficially this species could be con-
fused with P. salacinifera (Hale) Hale, which has a
pale brown lower surface and salazinic acid. The
two occupy very similar habitats (trees in open
secondary forests) and geographic ranges.
Specimens Examined. — United States: Florida, Hale 34139,
16859, 17668, 17696, Moore 3357 (DUKE, US), Rapp 311, 662
(US). Mexico: Veracruz, Hale 19389; Chiapas, Hale 19972
(DUKE, LD, US), Hale 20187 (BM, US), 20607a. Honduras: El
Paraiso, Standley 560 (F): Morazan, Standley and Williams
1643 (F). Cuba: Pinar del Rio, Imshaug 25289, 25310, 25333
(MSC); Oriente, Hioram 5963 (US). Puerto Rico: Howe 1162
(NY). Trinidad: Wedermann 2305 (8). Colombia: Santander,
Killip 15212, Nee and Mori 3768 (US). Venezuela: Merida,
Nash 2025 (US). Bolivia: El Beni, Wedermann 2305 (S). Brazil:
Minas Gerais, Vainio 588 (BM, TUR), 603 (BM, TUR); Rio de
Janeiro Burchell 946 (BM); Mato Grosso, Malme s.n., 20506b
(S), 2369, 2408 (LD, S, US). Guinea: N’Zerekore, Santesson
10573a (UPS). Taiwan: Asahina (TNS1.
Pseudoparmelia amplexa
Figure 6 d
Pseudoparmelia amplexa (Stirton) Hale, 1974:189.
Parmelia amplexa Stirton, 1877:212 [type collection: Somer-
set East, Union of South Africa, MacOwan (BM, lectotype;
GLAM, isolectotype)].
Parmelia caperata var. glaucopis Muller Argoviensis, 1894:258
[type collection: Matangiri, Seen Region, Africa, Stuhlmann
359 (559 on type label) (G, lectotype; BM, isolectotype)].
Parmelia glaucopis (Muller Argoviensis) Vainio, 1900:4.
Parmelia subconspersa var. benguellensis Vainio in Welwitsch,
1901:401 [type collection: Mt. Morro de Lopollo, Huilla,
Angola, Welwitsch 31 (BM, lectotype)].
Parmelia benguellensis (Vainio) Dodge, 1959:70.
Pseudoparmelia benguellensis (Vainio) Hale, 1974:189.
Description.— Thallus rather closely adnate on
bark, coriaceous, greenish yellow, 3-5 cm broad;
lobes sublinear, crowded, partly black rimmed, 1-2
mm wide; upper surface plane, dull, rugose and
becoming lobulate with age and heavily pycnidiate;
lower surface black and minutely velvety to the
margin, sparcely to moderately rhizinate. Apothecia
numerous, sessile, 1-2 mm in diameter; spores 8,
7-8 X 16-18 ju,m.
Chemistry.— Cortex K — , medulla K — , C — ,
KC — , or KC+ rose, P+ red; usnic acid and proto-
cetraric acid with or without a trace of atranorin.
Distribution. — Southern Africa.
Habitat. — On trunks and branches of trees in
open rocky areas.
Remarks. — I had at first identified all African
specimens as Pseudoparmelia rutidota (Hooker and
Taylor) Hale, but close examination showed that
they have consistently narrow lobes and a jet black
velvety lower surface (Figure la, b). By compari-
son, P. rutidota, which is known from Australia and
the New World, often has quite broad lobes (to
5 mm wide) and a shiny surface below and brown
bare zone near the tips. The sorediate morph of
P. amplexa is P. subamplexa Hale and the isidiate
morph is P. pachydactyla (Hale) Hale. All three
appear to lack any fatty acids, in contrast to the
P. rutidota group, which usually produces one or
more of the common fatty acids.
Specimens Examined. — Rhodesia: Kofler (LD, US), Schiitte
(LD, US), Sheppard 2 (US). Angola: Bie, Gossweiler 3256e
(BM, US), Degelius (Degelius herbarium, US); Huila, De-
gelius (Degelius herbarium); Moxico, Degelius (Degelius
herbarium, US).
NUMBER 31
17
Pseudoparmelia annexa
Figure 6e
Pseudoparmelia annexa (Kurokawa) Hale, 1974:189.
Parmelia annexa Kurokawa, in Hale and Kurokawa, 1964:151
[type collection: Caledon, Cape Province, Union of South
Africa, Almborn 5683 (LD, holotype; isotype in US)].
Description. — Thallus tightly adnate on rock,
ashy mineral gray, 3-7 cm broad; lobes subirregular
to sublinear, 0.7-8. 5 mm wide; upper surface plane,
shiny, continuous or irregularly cracked with age,
moderately isidiate, the isidia mostly simple, about
0.2 mm high, darkening at the tips; lower surface
black, sparsely rhizinate. Apothecia adnate, 1.5-5
mm in diameter, the amphithecium isidiate; spores
8, 5 X 7—8 pm.
Chemistry. — Cortex K+ yellow, medulla K — ,
C + , KC + red, P — ; atranorin and lecanoric acid.
Distribution. — Uganda, Angola, and Union of
South Africa.
Habitat. — On exposed rock outcrops from sea
level to 1200 m.
Remarks. — This rare saxicolous species is the
isidiate morph or close relative of P. molybdiza
(Nylander) Hale, which lacks isidia and has gen-
erally broader lobes. Both are African endemics.
Specimens Examined.— Uganda: Pian, Swinscow 6/1970
(BM, US), Other records from Angola and South Africa are
listed in Hale and Kurokawa (1964:151).
Pseudoparmelia aptata
Figure 6/
Pseudoparmelia aptata (Krempelhuber) Hale, 1974:189.
Parmelia aptata Krempelhuber in Nylander, 1869:291 [type
collection: Kotgurh, Himalayas, India, Skoliczka 509 (M,
lectotype; BM, H, K, US, isolectotypes)].
Parmelia asmarana Vainio, 1898b:37 [type collection: Asmara,
Ethiopia, Levander (TUR, Vainio herbarium number
2664, lectotype)].
Parmelia nipponica Zahlbruckner, 1927:353 [type collection:
Hachinoche, Japan, Faurie 1239 (W, lectotype)].
Description. — Thallus adnate on bark, whitish
mineral gray, 6-12 cm broad; lobes sublinear to
subirregular, more or less apically rotund, 2-5 mm
wide; upper surface plane, continuous or cracked
with age, laminally sorediate, the soralia capitate;
medulla white except for pale orange pigmentation
below the soralia; lower surface black, sparsely
rhizinate except for a narrow marginal papillate
zone. Apothecia not seen.
Chemistry.- — Cortex K+ yellow, medulla K — ,
C — , KC+ rose, P — ; atranorin, perlatolic acid, and
an unidentified pigment.
Distribution. — Africa, India, Indonesia, Aus-
tralia, and Japan.
Habitat. — On trees (Pinus, deciduous trees),
secondary forests at low to mid elevations.
Remarks. — This Old World lichen is externally
similar to P. texana (Tuckerman) Hale, which con-
tains divaricatic acid and is pantropical-pantem-
perate. It apparently evolved in the Old World as
a chemical variant from a now extinct fertile per-
latolic acid-containing progenitor, related perhaps
to P. nairobiensis (Steiner and Zahlbruckner) Hale.
It is basically sympatric with P. texana in most of
Asia and in Japan comprises about two-thirds of
the specimens in this group. Chemical tests must
be made to separate the species.
Specimens Examined. — Tanzania: Hoeg (TRH). Union of
South Africa: Natal, Almborn 6548, 8538, 8697, 9643 (LD),
Hoeg (TRH). India: Sikkim, Hara (TNS). Japan: Province
Aki, Hale 29517, 29542; Province Ohmi, Hale 29473. Indo-
nesia: Java, Groenhart 8853, 8854 (BOR), 2197, 5322, 5324
(L, US), Neeruoort 753 (BOR). Australia: Queensland, Bailey
(BM).
Pseudoparmelia arcana
Figure 7 a
Pseudoparmelia arcana (Kurokawa) Hale, 1974:189.
Parmelia arcana Kurokawa in Hale and Kurokawa, 1964:151
[type collection: Baba, Mocamedes, Angola, Degelius (De-
gelius herbarium: isotype in US)].
Description. — Thallus tightly appressed on rock,
ashy white, 3-7 cm in diameter; lobes sublinear,
crowded and appearing subareolate toward the
center, 0.3-1 mm wide; upper surface plane to con-
vex, continuous to tangentially cracked, moderately
isidiate, the isidia mostly simple, about 0.2 mm
high; medulla white or in part pale yellow; lower
surface brown to pale brown, sparsely rhizinate.
Apothecia not seen.
Chemistry. — Cortex K+ yellow, medulla K — ,
C — , KC — , P — ; atranorin, fatty acids, and un-
identified pigments.
Distribution. — Angola and Union of South
Africa.
18
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Habitat.— On rocks in dry open areas at about
1000 m elevation.
Remarks. — This rarely collected species seems to
be related to P. annexa (Kurokawa) Hale, which
contains lecanoric acid. The yellow pigmentation
does not seem to be a consistent character to judge
from the few specimens available.
Specimens Examined. — Union of South Africa: Basutoland,
Kofler (LD); Cape Province, Ha eg (TRH); Orange Free
State, Plank (PRE).
Pseudoparmelia baltimorensis
Ficure 7 b
Pseudoparmelia baltimorensis (Gyelnik and Foriss) Hale,
1974:189.
Parmelia baltimorensis Gyelnik and Foriss in Gyelnik, 1931a:
167 [type collection: Gunpowder River, Baltimore County,
Maryland, Plitt (BP, holotype; US, isotype)].
Description. — Thallus closely to loosely adnate
on rocks, yellowish green, up to 15 cm broad but
colonies often coalescing and covering extensive
areas of rock; lobes sublinear to irregular, apically
subrotund, 3-5 mm wide; upper surface plane, be-
coming covered toward the center with large, thick,
simple or branched isidioid pustules, fragile and
breaking open apically but not becoming sorediate;
lower surface black except for a narrow, smooth
to rugose, papillate, brown zone along the margins,
moderately rhizinate. Apothecia rare, substipitate,
1-3 mm in diameter, the amphitliecium pustulate;
spores 8, 6-7 X 13-15 pm.
Chemistry. — Cortex K — , medulla K — , C— or
C T rose, KC+ rose, P+ red; usnic acid, proto-
cetraric acid, and very frequently gyrophoric acid.
Distribution. — Eastern United States and ad-
jacent Canada.
Habitat. — Sandstone and granite rocks in open
deciduous forests.
Remarks. — This species had been confused with
the equally common but usually corticolous P.
caperata (L.) Hale. The distinction between the
coarse laminal pustules of P. baltimorensis and the
more or less diffuse soralia of P. caperata, as well
as the presence of gyrophoric acid as an accessory
substance in P. baltimorensis, was recently dis-
covered by Dr. W. L. Culberson (in litt.), who
kindly showed me his preliminary results. He is
now preparing a paper on the biology of these two
species that will present much more detailed in-
formation.
Representative Specimens Examined (all in US). — Canada:
Ontario, Hale. United States: Massachusetts, Hutchinson
1744; Connecticut, Hale 19076; New Jersey, Hale 17266;
Pennsylvania, Becking 57070315, Hale 16292, Heller, White
210; Maryland, Hermann 13768, Leonard 2933, 3086, Nor den
27, Plitt; West Virginia, Hale 10734, 13016; Kentucky, Allen
545, Hale 13115; Illinois, Hale 13926, 14029, Skorepa 943;
Wisconsin, Hale 23542, Thomson 919; Minnesota, Hale 23479;
Missouri, Greenman 4763, Ireland 3238, Schoop 254; Virginia,
Hale 15107, 17317, Luttrell 120, 4876; North Carolina, Cul-
berson in Lichenes Selecti Exsiccati 915, Hale 16371; Tennes-
see, Degelius, Hale 31118, Phillips 321; South Carolina, Hale
7747; Georgia, Hale 30877; Alabama, Hale 7125, 33841, Mc-
Cullough 344, 2032; Mississippi, Hale 7846; Arkansas, Hale
in Lichenes Americani Exsiccati 2, Hale 3644, 3813, Moore
259; South Dakota, Vischer 4; Kansas, Hale 2934, 4534; Okla-
homa, Hale 5053; Texas, Hale 34317. A number of records
in other herbaria would have been identified as Parmelia
caperata by me before 1970; it was impractical to borrow
these specimens again and correct the names.
Pseudoparmelia basutoensis
Figure 7c
Pseudoparmelia basutoensis (Hale) Hale, 1974:189.
Parmelia basutoensis Hale, 1972b:342 [type collection: Roma,
Basutoland, Union of South Africa, Kofler (LD, holotype;
isotype in US)].
Description. — Thallus adnate on rocks, pale
greenish white, 3-5 cm broad; lobes sublinear to
subirregular, contiguous, 2-3 mm wide; upper sur-
face plane, isidiate, the isidia cylindrical, simple;
lower surface pale brown, sparsely rhizinate. Apo-
thecia sparse, 1-3 mm in diameter, spores not found.
Chemistry. — Cortex K+ yellow, medulla nega-
tive with all reagents; atranorin and an unknown
substance.
Distribution. — Union of South Africa.
Habitat. — Rocks in dry regions.
Remarks. — As I pointed out in the original de-
scription, this species has the general aspect of a
Xanthoparmelia but lacks usnic acid. The non-
isidiate morph and presumptive parent is P. prolata
(Hale) Hale (see below), but neither species has
been collected in sufficient quantity for more de-
finitive statements on biology and possible evolu-
tion of the group. The unknown substance appears
to be a new fatty acid that reacts faintly yellow with
sulfuric acid. The same substance has been dis-
covered in a brown Parmelia, P. scabrella Esslinger
(Esslinger, 1976).
NUMBER 31
19
Figure 7. — Species of Pseudoparmelia: a, P. arcana (Degelius, isotype in US); b, baltimoren-
sis (Luttrell 3369 in US); c, P. basutoensis (Hewitt, holotype in TRH); d, P. caperata (Plitt in
US); e, P. caribaea (Gallo 494, holotype in US); f, P. carneopruinata (Hale 42912). (Scale in mm.)
20
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Psendoparmelia callichroa Kurokawa, new species
Description. — Thallus adnatus, ramulicola,
viridi-flavicans, 3-6 cm latus, lobis subirregularibus,
congestis, apice subrotundatis; superne planus vel
mox irregulariter rugosus, isidiis sorediisque desti-
tutus; cortex superior 10-13 ^ra crassus, epicortex
perforatus, stratum gonidiale 12 jxm crassum, me-
dulla alba, 100-120 crassa, cortex inferior 12-13
/j,m crassus; subtus niger, nitidus, modice rhizinosus,
rhizinis simplicibus, nigris. Apothecia numerosa,
amphithecio ruguloso, margine crenato, 1. 0-4.5 mm
diametro; hymenium 70-85 ^m altum; sporae 8:nae,
6-8 X 13-16 /mi.
Chemistry. — Cortex K — , medulla negative with
all reagents; usnic acid and protolichesterinic acid.
Holotype. — Chile: Colchagua, Hacienda de Con-
quenes, P. Dusen 91, 22 August 1896 (S; US, iso-
type).
Distribution. — Chile.
Habitat. — On twigs.
Remarks. — This species would probably be
identified as P. rutidota (Hooker and Taylor) Hale
without a chemical test (P. rutidota is P+ red with
protocetraric acid). The thallus is smaller but we
need many more collections to determine whether
any morphological characters correlate with the
chemical difference.
Pseudoparmelia caperata
Figure Id
Pseudoparmelia caperata (L.) Hale, 1974:189.
Lichen caperatus L. 1753:1147 [type collection: England,
Dillenius, Historia Muscorum 193, specimen represented
by pi. 25: fig. 97 (OXF, lectotype)].
Parmelia caperata (L.) Acharius, 1803:216.
Parmelia caperata b. cyliphora Acharius, 1814:196 [type col-
lection: North America, Muhlenberg (H, lectotype; UPS,
isolectotype); listed in error as "cylisphora."]
Parmelia perlata var. flavicans Tuckerman, 1866:13 [type
collection: California, Bolander 70 (FH-Tuck, holotype)].
Parmelia euplecta Stirton, 1877-78:299 [type collection: Bris-
bane, Australia, Bailey (BM, lectotype)].
Parmelia flavicans (Tuckerman) Tuckerman, 1882:53 [not
P. flavicans (Swartz) Acharius, 1803:268 (= T eloschistes)].
Parmelia ochroleuca f. sorediosa Muller Argoviensis, 1883b:
77 [type collection: Mt. Dromedary, Australia, Reader (G,
lectotype)].
Parmelia subglauca Nylander in Gasilien, 1894:126 [type
collection: Saint Omer, France (H, lectotype: BM, PC iso-
types)].
Parmelia cylisphora [sic] (Acharius) Vainio, 1896b:7.
Parmelia caperata var. subglauca (Nylander in Gasilien) Ny-
lander, 1896:35.
Parmelia caperata f. laevissima Gyelnik, 1928:587 [type col-
lection: Meleghegy, Feher, Hungary, Gyelnik (BP, holo-
type)].
Parmelia herreana Zahlbruckner, 1929:239 [based on P. flavi-
cans (Tuckerman) Tuckerman],
Parmelia pseudosorediosa Gyelnik, 1931b:288 [based on Par-
melia ochroleuca f. sorediosa Muller Argoviensis].
Parmelia negativa Gyelnik 1934:301 [type collection: Salisbury
Cove, Maine, Plitt (BP, holotype)].
Pseudoparmelia euplecta (Stirton) Hale, 1974:190.
Pseudoparmelia pseudosorediosa (Gyelnik) Hale, 1974:190.
Description. — Thallus adnate to loosely attached,
growing on bark or more rarely on rocks, yellowish
green, 5-20 cm in diameter, coalescing to form
larger colonies; lobes subirregular, apically rotund,
3-8 mm wide; upper surface plane to undulate or
rugulose, continuous, sorediate, the soralia laminal,
diffuse and coalescing; medulla rarely orange near
the lower cortex; lower surface black, moderately to
sparsely rhizinate except for a naked brown rugose
zone along the margins. Apothecia very rare, 2-5
mm in diameter; spores 8, 8-13 X 17-24 /xm.
Chemistry. — Cortex K-, medulla K — , C — ,
KC+ rose, P-f- red, atranorin, caperatic, proto-
cetraric, and usnic acids; if pigmented, skyrin pres-
ent.
Distribution. — Pantemperate on all major conti-
nents (Figure 8).
Habitat. — On conifers and deciduous trees in
temperate forests, common on rocks only at the
northern part of its range, from sea level to over
3000 m elevation.
Remarks. — This is by far the most widespread
and commonly collected foliose lichen in all tem-
perate areas of the world. The range in morpho-
logical variation is very small and the chemistry
remarkably uniform. In eastern North America it
can be confused with the primarily saxicolous P.
baltimorensis (see discussion above). The presump-
tive nonsorediate parent seems to be P. rutidota
(Hooker and Taylor) Hale (see below).
Older authors consistently placed this species in
Parmelia section Amphigymnia (— Parmotrema ).
The lobes, however, are clearly adnate at the mar-
gins, the bare rim below is very narrow, the tips of
rhizines near the margin are frayed, and the apo-
thecia are adnate and nonperforate. This combina-
tion of characters relates the species much mdre
closely to Pseudoparmelia than to Parmotrema.
The species is apparently present in Australia
NUMBER 31
21
Figure 8.— World distribution of Pseudoparmelia caperata. (Dots may represent more than one
collection.)
and New Zealand, represented by the anonomously
pustulate P. euplecta. Parmelia ochroleuca f. sore-
diosa, also described from Australia, is atypical in
having rather discrete orbicular soralia. Future
studies in this region may show that these popula-
tions are distinct from P. caperata.
Specimens Examined. — See Figure 8 for localities of speci-
mens annotated in the major herbaria. Some specimens from
eastern North America which I annotated before about 1972,
especially those on rock, will be P. baltimorensis.
Pseudoparmelia caribaea
Figure le
Pseudoparmelia caribaea (Hale) Hale, 1974:189.
Parmelia caribaea Hale in Hale and Kurokawa, 1964:152
[type collection: St. Barthelemy, West Indies, Le Gallo 494
(US, holotype)].
Description. — Thallus adnate on rocks, ashy
white, rather coriaceous, 6-10 cm broad; lobes ir-
regularly sublinear, often densely imbricate, 3-6
mm wide; upper surface plane or rugulose, cracked
with age; lower surface black, moderately rhizinate.
Apothecia subpedicellate, at first globose, urceolate
at maturity, up to 5 mm in diameter; spores 8, 4-5
X 10-13 jx m.
Chemistry. — Cortex K+ yellow, medulla K — ,
C — , KC+ rose, P+ red; atranorin and protocetraric
acid.
Distribution. — West Indies and French Guiana.
Habitat. — On schist and volcanic rocks in open
areas up to 275 m elevation.
Remarks. — Except for the two specimens from
French Guiana and the Isle of Pines, P. caribaea
is confined to St. Barthelemy, where it has been col-
lected many times. There are no close relatives in
the genus.
Specimens Examined. — Cuba: Isle of Pines, Britton et al.
14436, 15237 (US). St. Barthelemy: Le Gallo 403, 424, 527,
536, 546 (US), 2604 (MSC, US). French Guyana: Degelius
(Degelius herbarium).
22
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Psendoparmelia carneopruinata
Figure 7/
Pseudoparmelia carneopruinata (Zahlbruckner) Hale, 1974:
189.
Parmelia carneopruinata Zahlbruckner, 1902:419 [type col-
lection: Rio de Janeiro, Brazil, Hohnel 164 (W, lectotype)].
Parmelia sbarbaronis Bouly de Lesdain, 1923:278 [type col-
lection: Catalupe, Varezze, Liguria, Italy, Gresino 11467 (F,
lectotype)] .
Description. — Thallus closely adnate, corti-
colous, greenish mineral gray or in the herbarium
buff, 5-9 cm in diameter; lobes sublinear, 1-2.5 mm
wide; upper surface strongly reticulately ridged and
rugulose, becoming pruinose near the lobe tips,
sometimes densely lobulate, sorediate, soralia about
1 mm wide, often coalescing; lower surface black,
sparsely rhizinate. Apothecia rare, adnate, 1-4 mm
in diameter, the disc pruinose or naked, the am-
phithecium sorediate; spores 8, 6-9 X 9-13 p.m.
Chemistry. — Cortex K+ yellow, medulla K +
yellow, C — , KC — , P+ pale orange; atranorin,
stictic acid, and constictic acid with associated un-
knowns.
Distribution. — Mexico, Central America, West
Indies, Colombia, Venezuela, Brazil, Uruguay, Ar-
gentina, and southern Europe.
Habitat. — On shade trees in banana and coffee
plantations and on deciduous trees in open pastures
at 300-2000 m elevation.
Remarks. — This species is closely related to P.
scrobicularis (Krempelhuber) Hale, which is or at
least may be the nonsorediate progenitor. Another
very close sorediate species, P. crozalsiana (Bouly de
Lesdain) Hale, has much broader, subrotund lobes
and a wider, more temperate distribution. The two
species, however, probably intergrade and are not
always easily distinguished.
Specimens Examined. — Mexico: Nayarit, Weber 33609
(COLO); Veracruz, Hale 19398, 19432 (DUKE, US), 19452,
19654, 19672, 19794, 21081, 21195. Honduras: Morazdn,
Standley 11655 (F, US), 11657 (F). Costa Rica: San Jose,
Standley 34811 (US). Jamaica: Orcutt 5603 (US), Plitt (US).
Colombia: Lindig 2590 (G). Venezuela: Merida, Hale 42785,
42912. Brazil: Minas Gerais, Mexia 5322 (BM); Sao Paulo,
Schindler 4512, 4525 (KR, US). Uruguay: Canelones, Osorio
6236 (MVM). Argentina: Jujuy, Fries 43b (S). Europe: Italy,
Sbarbaro (TUR).
Pseudoparmelia caroliniana
Figure 9 a
Pseudoparmelia caroliniana (Nylander) Hale, 1974:189.
Parmelia caroliniana Nylander, 1885:614 [type collection:
South Carolina, Ravenel 404 (H, lectotype; FH-Tuck, iso-
lectotype)].
Parmelia perlata var. subrevoluta Muller Argoviensis, 1880:267
[type collection: Petropolis, Brazil, Deventer 45 (G, lecto-
type)].
Parmelia isidiophora Zahlbruckner, 1902:420 [type collection:
Botanical Garden, Rio de Janeiro, Brazil, Hohnel 169, (W,
lectotype)].
Parmelia luteola Zahlbruckner, 1909:170 [type collection:
Near Barra Mansa, Sao Paulo, Brazil, Wettstein and Schiff-
ner (W, lectotype)].
Parmelia wainoana Lynge, 1914:87 [type collection: Santa
Anna da Chapada, Mato Grosso, Brazil, Malme 2435C (S,
lectotype)].
Pyxine azorea Nylander, 1895:100 [type collection: Azores,
Michel 66 (H, lectotype)].
Description. — Thallus closely adnate to bark,
rarely on rock, 5-10 cm broad; lobes subirregular,
rotund, 2-5 mm wide; upper surface plane, re-
ticulately white-maculate and finely cracked, densely
isidiate, the isidia 1-3 mm high, often branched;
lower surface black, very rarely uniformly dark
brown, moderately rhizinate except for a narrow
naked or papillate zone along the margins. Apo-
thecia rare, adnate, 1-3 mm in diameter, the am-
phithecium isidiate; spores 8, 6-8 X 13-15 p.m.
Chemistry. — Cortex K+ yellow, medulla K — ,
C — , KC+ faint rose or purple, P — ; atranorin and
perlatolic acid with associated unknowns.
Distribution. — Southeastern United States,
Mexico, Central America, West Indies, Venezuela,
Ecuador, Brazil, Azores, Africa, and Thailand.
Habitat. — On trees (conifers and deciduous
trees) in coffee plantations, secondary forest, and
pastures and rarely on rocks from sea level to 2800
m elevation.
Remarks. — The first study of this species was
done by Culberson (1957), who clarified its re-
lation and frequent confusion with Parmelia
rudecta Acharius, another widespread species which
differs in having pseudocyphellae and lecanoric acid
as well as a paraplectenchymatous upper cortex.
Pseudoparmelia caroliniana is the only isidiate spe-
cies in the genus with perlatolic acid; P. aptata
(Krempelhuber) Hale also has perlatolic acid but
is sorediate and lacks the fine reticulation so charac-
teristic of P. caroliniana. I wish to thank Dr. T. D.
V. Swinscow for pointing out the synoymy of
Pyxine azorea.
Specimens Examined. — United States: See Culberson (1957)
for localities over the range in the United States and Moore
(1968:223) for specimens from Florida. Mexico: Hidalgo,
NUMBER 31
23
Figure 9. — Species of Pseudoparmelia: a, P. caroliana (Hale 20144); b, P. chapadensis (Malme
2297B, lectotype in S); c, P. cinerascens ( Xavier 713 in US); d, P. concomitans ( Hill 12091, holo-
type in BM); e, P. concrescens (Sw in scow 2U 24/21 in BM); /, P. condyloides (Almborn 4882,
holotype in LD). (Scale in mm.)
24
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Robinson (US); Veracruz, Hale 19471, 19488, 21131, 21135;
Chiapas, Hale 19900, 20104, 20144, 20215, 21310, 20333. Hon-
duras: Morazan, Standley 238, 11603 (F). Nicaragua: Standley
8432 (F). Costa Rica: Guanacaste, Standley 44370 (US).
Panama: Chiriqui, Hale 38804. Cuba: Wright 71 (FH, M, PC,
UPS, US); Oriente, Imshaug 24763, 24858 (MSC). Jamaica:
Imshaug 13762, 14200 (MSC), Plitt (US). Haiti: Ouest, Imshaug
22542, 22859, 22954, 23061 (MSC, US), Wetmore 2882, 3076,
3149, 3207 (MSC, US), Leonard 3709b (US). Dominican Re-
public: Cordillera Central, Imshaug 23714 (MSC, US, WIS),
23718 (BM, G, MSC); La Vega, Allard 18033 (US). Venezuela:
Distrito Federal, Dennis 1503 (BM). Santesson 6672, 6681 (S);
Merida, Hale 42039, 42196, 43318, 44081, 44225; Tdchira,
Hale 45632, 45649. Ecuador: Galapagos, Johannsen 10 (WIS),
Weber 40031 (COLO, UPS). Brazil: Rio de Janeiro, Eiten
7373 (US); Sao Paulo, Eiten 2978, (US) Osorio 4925 (MVM);
Santa Catarina, Reitz and Klein 13145, 15398 (US). Europe:
Portugal, Persson (UPS). Ivory Coast: Man, Santesson 10494
(UPS, US). Uganda: Burahiya County, Swinscow 2U 18 (BM);
Kyadondo County, Swinscow 2U 1/7 (BM). Angola: Huila,
Degelius (Degelius herbarium). Laos: Tsuyama 19 (TNS).
Pseudoparmelia chapadensis
Figure 95
Pseudoparmelia chapadensis (Lynge) Hale, 1974:189.
Parmelia chapadensis Lynge, 1914:153 [type collection: near
Bocca da Serra, Serra da Chapada, Mato Grosso, Brazil,
Malme 2297B (S, lectotype)].
Description. — Thallus closely adnate on rock,
yellowish green, 1-1.5 cm broad; lobes sublinear,
short, 0.5 mm wide, convex and expanded at the
tips; medulla yellowish; lower surface pale brown,
moderately rhizinate. Apothecia common, sessile,
0. 5-1.0 mm in diameter; spores 8, 4-5 X 8-10 pm.
Chemistry. — Cortex K+ yellow, medulla K + ,
C + , KC-f- yellowish, P+ red; unidentified sub-
stances present.
Distribution. — Brazil.
Habitat. — Rocks in open areas.
Remarks. — This peculiar species is tentatively
placed in Pseudoparmelia because the lobes are
eciliate and the rhizines simple. Contrary to my
earlier findings (Hale, 1960), it does not contain
usnic acid or protocetraric acid. The yellow colora-
tion is caused by unidentified pigments in both the
cortex and the medulla. Perhaps additional collec-
tions will clarify the status of the species.
Pseudoparmelia cinerascens
Figure 9c
Pseudoparmelia cinerascens (Lynge) Hale, 1974:189.
Parmelia cinerascens Lynge, 1914:104 [type collection: Para-
guari, Paraguay, Malme 1498 (S, lectotype)].
Description. — Thallus adnate, fragile, ashy min-
eral gray, 4-5 cm broad; lobes short, becoming
imbricate, 1-S mm wide; upper surface plane, ir-
regularly cracked with age, densely isidiate, isidia
cylindrical, simple or branched; lower surface black,
rhizines moderate to sparse. Apothecia abundant,
adnate; spores 8, 6 X 12 pm.
Chemistry. — Cortex K+ yellow, medulla K-f-
yellow turning red, C — , KC — , P+ orange; at-
ranorin and salazinic acid.
Distribution. — Brazil and Paraguay.
Habitat. — On trees in open forest.
Remarks. — This rare species is superficially near
P. salacinifera (Hale) Hale, which differs in having
a pale brown lower surface.
Specimens Examined. — Brazil: Pernambuco, Xavier 713 (US).
Pseudoparmelia concomitans, new species
Figure 9 d
Description. — Thallus laxe adnatus, corticola,
bubalino-albidus, coriaceus, 4-6 cm latus, lobis sub-
irregularibus, apice rotundatis, 3-5 mm latis;
superne planus, nitidus albo-maculatusque, sorediis
et isidiis destitutus; cortex superior 10-14 pm cras-
sus, epicortex sparse perforatus, stratum gonidiale
10-12 pm crassum, medulla 90-110 pm crassa.
Cortex inferior 8-10 pm crassus; subtus niger,
sparse rhizinosus, ambitu nudus, castaneus, nitidus.
Apothecia numerosa, substipitata, 1-2 mm dia-
metro; sporae 8:nae, 12 X 20-23 pm.
Chemistry. — Cortex K+ yellow, medulla K — ,
C + , KC+ red, P+ red, atranorin, gyrophoric acid,
and protocetraric acid.
Holotype. — New Caledonia: Baie des Crabes, lie
de Pins, on fallen branches of Araucaria cookii, D. J.
Hill 12124 (BM; US, isotype) .
Distribution. — New Caledonia.
Habitat. — On branches of Araucaria.
Remarks. — This species is known only from the
type collection and seems unrelated to any other
Pseudoparmeliae. The lobes are coriaceous and
strongly white maculate. Only P. martinicana pro-
duces this combination of medullary substances but
it is an isidiate, fragile lichen confined to the West
Indies.
Pseudoparmelia concrescens
Figure 9e
Pseudoparmelia concrescens (Vainio) Hale
Parmelia concrescens Vainio, 1901:400 [type collection: Serra
NUMBER 31
25
da Xella, Huila, Angola, Welwitsch 30 pro parte (TUR,
lectotype; BM, isolectotype)].
Parmelia capensis Nylander, 1885:613 [type collection: Cape
of Good Hope, Union of South Africa, Drege (H, Nylander
herbarium number 35174, lectotype; PC, isotype); not
P. capensis (Acharius) Sprengel, 1827:280 (= Teloschistes)].
Parmelia austroajricana Zahlbruckner, 1929:152 [type: based
on P. capensis NylanderJ.
Parmelia caffrorum Zahlbruckner, 1932a:555 [type: based on
P. capensis Nylander].
Description. — Thallus adnate on bark, ashy
mineral gray, 4-10 cm in diameter; lobes irregularly
sublinear, 1-3 mm wide; upper surface plane, con-
tinuous or cracked on older lobes, densely isidiate,
the isidia short, cylindrical, mostly simple; lower
surface black, sparsely rhizinate. Apothecia rare,
adnate, 2-3 mm in diameter; spores 8, 5 X 7-8 ^m.
Chemistry. — Cortex K+ yellow, medulla K — ,
C — , KC+ faint wine red, P — ; atranorin and di-
varicatic acid with associated unknowns.
Distribution. — Southern Africa.
Habitat. — On trees or rocks in open woodland
at 1000-2300 m elevation.
Remarks. — This is one of the commoner, more
widespread lichens in Africa. It is presumed to be
an isidiate morph of P. nairobiensis (Steiner and
Zahlbruckner) Hale, with which it is sympatric.
It is also close to the more widespread P. ecaperata
(Muller Argoviensis) Hale, which differs in con-
taining usnic acid in the cortex.
Specimens Examined. — Uganda: Dummer 602 (US); Busiro
County, Swinscow 2U 24/21 (BM, US); Kinkizi County, Dale
L46 (BM, US). Zaire (Congo): Degelius (Degelius herbarium,
US), Hoeg (TRH), Louis 7494 (BR), de Witte 2675 (BR).
Angola: Cuanza-Sul, Degelius; Bie, Degelius; Huila, Degelius;
Moxico, Degelius (all Degelius herbarium). Rhodesia: Arnell
1282a (LD), Schiitte 46c (LD, US). Mozambique: Almborn
6888, 7015 (LD). Union of South Africa: Natal, Almborn
8637 (LD).
Pseudoparmelia condyloides
Figure 9/
Pseudoparmelia condyloides (Kurokawa) Hale, 1974:189.
Parmelia condyloides Kurokawa in Hale, 1972:343 [type col-
lection: Kamieskroon, Namaqualand, Union of South
Africa, Almborn 4882 (LD, holotype; TNS, US, isotypes)].
Description. — Thallus closely adnate on rock,
dark mineral gray with a yellowish cast, 3-5 cm
broad; lobes subirregular to sublinear, crowded,
1.5-2. 5 mm wide; upper surface plane to rugose-
bullate, the cortex breaking away easily; lower sur-
face pale brown, moderately rhizinate, the rhizines
pale. Apothecia numerous, adnate, 1-2 mm in di-
ameter; spores 8, 4 X 5 pm.
Chemistry. — Cortex K+ yellow, medulla K +
yellow turning red, C — , KC — , P-f orange; atra-
norin, “chalybeizans” unknown, salazinic acid, and
a pale yellow unidentified pigment.
Distribution. — South Africa.
Habitat. — On rocks in open dry areas.
Remarks. — This is another species of Pseudopar-
melia that bears a close resemblance to Xanthopar-
melia. The “chalybeizans” unknown, in fact, was
previously known only in Xanthoparmelia (Hale,
1972b) . This specimen apparently endemic to the
arid regions of South Africa.
Specimens Examined. — Material from the Union of South
Africa is listed in Hale (1972b:343).
Pseudoparmelia conlabrosa, new species
Figure 10a
Description. — Thallus arete adnatus, corticola,
viridi-albidus, 3-5 cm latus, lobis sublinearibus,
apice plus minusve subrotundatis, congestis, 2-2.5
mm latis; superne undulatus, nitidus, apice albo-
reticulatus, dense isidiatus, isidiis cylindricis, pro
maxima parte simplicibus, usque ad 0.5 mm aids;
cortex superior 10-12 p m crassus, epicorticatus, epi-
cortice perforato, stratum gonidiale ca 10 pm
crassum, medulla alba, ca 100 pm crassa, cortex in-
ferior paraplectenchymatus, 8-10 pm crassus; subtus
niger, modice rhizinosus, rhizinis nigris, simplici-
bus. Apothecia ignota.
Chemistry. — Cortex K+ yellow, medulla K — ,
C T , KC+ red, P— ; atranorin and lecanoric acid.
Holotype. — Australia: New South Wales, be-
tween Majors Creek and Araluen, Weber and
McVean L-47102, 18 October 1967 (US; COLO,
isotype).
Remarks. — This species is obviously related to
Pseudoparmelia labrosa (Zahlbruckner) Hale, a
sorediate species with lecanoric acid also known
from Australia. No parent morph has yet been dis-
covered for these two species.
Pseudoparmelia corrugativa, new combination
Parmelia corrugativa Kurokawa and Filson, 1975:38 [type
collection: Near Bulhunnah, South Australia, Rogers 553
(MEL, holotype) (not seen)].
26
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Figure 10. — Species of Pseudoparmelia: a, P. conlabrosa (Weber and McVean L-47102, holo-
type in US); b, P. crozalsiana ( Culberson 6586 in US); c, P. cryptochlorophaea (Hale 19771);
d, P. cyphellata (Malme 2532B, isolectotype in US); e, P. dahlii (Dahl, isotype in US); f, P.
ecaperata (Tsuyama 5 in US). (Scale in mm.)
NUMBER 31
27
Description. — Thallus closely adnate on twigs,
whitish mineral gray, about 6 cm broad; lobes
irregularly elongate, apically rotund, 1.5-3 mm
wide; upper surface rugulose; medulla white in the
upper part and deep yellow in the lower half; lower
surface black, sparsely rhizinate. Apothecia numer-
ous, 1-4 mm in diameter; spores 8, 7 X 10 pm.
Chemistry. — Cortex K+ yellow, medulla nega-
tive with color reagents except for the pigment
which is K+ purple, atranorin, and unidentified
substances.
Distribution. — South Australia.
Habitat. — On twigs of tree in open area.
Remarks. — This species is very similar to P.
subtiliacea (Nylander) Hale, which differs in hav-
ing a white medulla without pigments but with
fatty acids. One additional specimen, which I have
not seen, was collected by Rogers (95) in Para-
Wirra National Park, South Australia. The species
is well illustrated in Kurokawa and Filson (1975)
(pi. 1: fig. 3).
Pseudoparmelia crozalsiana
Figure 10 b
Pseudoparmelia crozalsiana (Bouly de Lesdain) Hale, 1974:
189.
Parmelia crozalsiana Bouly de Lesdain ex Harmand, 1909:
555 [type collection: Agde, Herault, France, De Crozals,
May 1909 (US, lectotype)].
Description. — Thallus adnate to bark, greenish
mineral gray, 5-10 cm broad; lobes subirregular, 3-6
mm wide; upper surface strongly reticulately ridged
and wrinkled, sometimes white-pruinose, sorediate,
the soralia often produced along the ridges, coa-
lescing; lower surface black, moderately rhizinate
except for a narrow brown marginal zone. Apo-
thecia not seen.
Chemistry. — Cortex K+ yellow, medulla K +
yellow, C — , KC — , P+ pale orange, atranorin,
stictic acid, constictic acid, and associated un-
knowns.
Distribution. — Eastern United States, Mexico,
Brazil, Uruguay, Argentina, France, Italy, Zaire,
Union of South Africa, and India.
Habitat. — On trees in open secondary forests
at 100-2000 m elevation.
Remarks. — I first identified this species in North
America in 1960 (Hale, 1960) where it had been
misidentified as Parmelina aurulenta (Tuckerman)
Hale, a ciliate species. Since then it has been recog-
nized from many localities. The lobes are usually
quite broad and apically subrotund and the reticu-
late wrinkles very distinct without magnification.
Once the chemistry is recognized, it could only be
confused with Parmelia carneopruinata (see be-
low), which has lobes less than half as wide on
the average. There seems to be no nonsorediate
progenitor extant, unless we consider it to be the
rare African species P. inhaminensis (Dodge) Hale.
When I first examined the surface of this species
with the scanning-electron microscope (Hale,
1973a), I noted a strongly nodular surface without
pores, an anamolous condition for the genus. How-
ever, examination of other species has shown that
pores do occur rarely but that even though the
species does not seem to have a paraplectenchy-
matus upper cortex, as one would expect in a
nonepicorticate Parmelia, the cortical structure of
this species is rather atypical.
Specimens Examined. — United States: Illinois, Hale 13980,
Skorepa 1757 (US); Indiana, Hale 14097; Kentucky, Hale
13691; Ohio, Hale 13576; Virginia, Hale 15232, 15761, 18455;
Arkansas, Keck 1216 (US); Alabama, Hale 33829. Mexico:
Mexico, Hinton 7724 bis (BM); Veracruz, Hale 19494. Venez-
uela: Merida, Hale 42288, 42312, 43049a; Tdchira, Hale
45704. Brazil: Mato Grosso, Malme 2243C (S); Parand, Montes
10121 (DUKE); Pernambuco, Xavier 700 (US). Uruguay:
Lavalleja, Osorio 3667 (DUKE); Maldonado, Osorio 4879
(US); Rivera, Osorio 1075 (US). Argentina: Tucuman, Ven-
turi 336 (DAR). Europe: Italy, Davies (BM), Sbarbaro (US),
Sbarbaro in Lichenotheca Parva 32 (M), Sbarbaro in Lichenes
Selecti Exsiccati 96 (H, LD, US). Uganda: Mawakota County,
Swinscow 2U 41/2 (BM, US). Zaire: Louis 8150 (BR). Union
of South Africa: Natal, Almborn 8636 (LD), Hoeg (TRH).
India: Tamil Nadu, Awasthi 4276 (Awasthi herbarium), Hale
43976.
Pseudoparmelia cryptochlorophaea
Figure 10c
Pseudoparmelia cryptochlorophaea (Hale) Hale, 1974:189.
Parmelia cryptochlorophaea Hale, 1959:18 [type collection:
Ciudad Trujillo, Dominican Republic, Allard 15715a (US,
holotype)].
Description. — Thallus adnate to appressed on
large branches, 5-10 cm in diameter, light mineral
gray but soon turning buff in the herbarium; lobes
3-5 mm wide, apically subrotund, crowded; upper
surface rugulose and shiny, more or less reticulate
white-maculate at the tips, becoming cracked in
older parts; margins sorediate, the soredia in dense
28
SMITHSONIAN CONTRIBUTIONS TO BOTANY
erect soralia, 0. 6-1.0 mm thick, 1-1.5 mm high;
lower surface black and rhizinate at the center,
brown and naked to papillose in a narrow zone at
the margins. Apothecia rare, 1-3 mm in diameter,
spores 8, 4-5 X 6-9 pm.
Chemistry. — Cortex K+ yellow, medulla K — ,
C — , KC+ rose, P — ; atranorin, caperatic acid (C.
Culberson, 1965) , and cryptochlorophaeic acid.
Distribution. — Southern United States, Mexico,
Central America, West Indies, Venezuela, and
Brazil.
Habitat. — On trees (cashew, Xylopia, bamboo,
etc.) in mature forests at sea level to 500 m eleva-
tion.
Remarks. — The most unusual feature of this
New World species is the chemistry, the only re-
ported occurrence of cryptochlorophaeic acid in the
Parmeliaceae. The capitate soralia are also unique.
The white reticulation is similar to that in the
Parmelia texana group, with which it is obviously
closely allied.
Specimens Examined. — United States: Georgia, Hale 16797,
21985; Florida, Hale 16989, 17584, 17609, 21674, 21815, 21839,
Rapp (US) (for further records from Florida in DUKE see
Moore, 1968:225). Mexico: Veracruz, Hale 19771, 19811. Hon-
duras: Comayagua, Standley 5462, 5930 (F). Jamaica: Imshaug
15967, 15969 (US), Plitt s.n. (US). Dominican Republic: La
Vega, Allard 16833, 16845, 16854, 16861, 18028; Santiago,
Imshaug 23884 (MSC, US); Santo Domingo, Allard 15715,
16182, 16187, 16191a (US). Trinidad: Degelius s.n. (Degelius
herbarium); Lassen (C); Lewis 347 (PH). Venezuela: Barinas,
Steyermark and Rate 96533 (US); Merida, Hale 42505a.
Brazil: Ceara, Cutler 8072c (F); Rio de Janeiro, Glaziou 1848
(M).
Pseudoparmelia cyphellata
Figure 10 d
Pseudoparmelia cyphellata Lynge, 1914:15 [type collection:
Santa Anna da Chapada, Mato Grosso, Brazil, Malme
2532B (S, lectotype; LD, UPS, US, isolecto types)].
Parmelia cyphellata (Lynge) Santesson, 1942:473.
Description. — Thallus as in P. sphaerospora (see
below) except the upper surface isidiate, isidia
cylindrical, rather sparse, up to 1 mm tall. Apo-
thecia abundant, adnate to substipitate, 2-3 mm
in diameter; spores 8, about 5X7 pm.
Chemistry. — Cortex K+ yellowish, medulla yel-
lowish to orange with color reagents; atranorin,
stictic acid, and an unidentified pigment.
Distribution. — Brazil.
Habitat. — On trunks of trees in forest.
Remarks. — This species is clearly the isidiate
morph of P. sphaerospora (Nylander) Hale. Un-
fortunately it is still known only from the type
locality.
Pseudoparmelia dahlii, new species
Figure lOe
Thallus adnatus, corticola, viridi-flavicans, 3-4
cm latus, lobis sublinearibus, contiguis, 1. 5-2.0
mm latis; superne convexus, opacus, dense isidiatus,
isidiis cylindricis, simplicibus, usque ad 0.5 mm altis;
cortex superior 12-14 pm crassus, epicorticatus;
stratum gonidiale 15-20 pm crassum; medulla alba,
140-160 pm crassa; cortex inferior paraplectenchy-
matus, 12-14 pm crassus; subtus pallide castaneus,
dense rhizinosus, rhizinis pallidis, simplicibus, elon-
gatis. Apothecia adnata, 1 mm diametro; hymenium
40-45 pm altum; sporae 8:nae, 6 X 7-9 pm.
Chemistry. — Cortex K — , medulla K — , C + ,
KC+ red, P — , usnic acid, caperatic acid and le-
canoric acid.
Holotype. — Sri Lanka: Polonnaruwa, E. Dahl, 6
January 1972 (O; US isotype).
Distribution. — Sri Lanka.
Habitat. — On trees in lowland forest.
Remarks. — Superficially this lichen resembles P.
malaccensis (Nylander) Hale very closely, but the
rhizines are denser and longer, the lobes thicker
and dull without alboreticulate patterning, and
the chemical components different. It is, however,
clearly a member of the P. intertexta-P. malaccensis
lowland rain forest complex. Another closely related
species, P. rahengensis (Vainio) Hale, which occurs
in the higher elevation monsoon forest in Thailand,
is also externally very close. Chemical tests are
needed to identify the species. I wish to thank Dr.
Hildur Krog for allowing me to see the specimens
and to describe them.
Specimens Examined. — Sri Lanka: Dambulla, Dahl (O, US).
Pseudoparmelia ecaperata
Figure 10/
Pseudoparmelia ecaperata (Muller Argoviensis) Hale, 1974:
190.
Parmelia ecaperata Muller Argoviensis, 1891:378 [type col-
lection: Shire River, “Zambesica,” Africa, Kirk (G, lecto-
type; BM, isolectotype)].
NUMBER 31
29
Parmelia malaccensis var. laeteflavens Vainio, 1921:38 [type-
collection: Doi Sutep, Thailand, Hosseus, 1904 (TUR,
Vainio herbarium number 2764, lectotype)].
Parmelia laeteflavens (Vainio) Gyelnik, 1938a:32.
Parmelia djalonensis des Abbayes, 1951:966 [type collection:
Fouta-Djalon, Dalaba, Guinea, des Abbayes (REN, lecto-
type)]
Description. — Thallus as in P. concrescens (see
above) except yellowish green. Apothecia rare, 1-4
mm in diameter, the amphithecium isidiate; spores
8, 6 X 11-14 ,ji m.
Chemistry. — Cortex K — , medulla K — , C — ,
KC+ faint wine red, P — ; atranorin, divaricatic
acid, and usnic acid, rarely with protolichesterinic
acid.
Distribution. — Africa, Nepal, India, and Thai-
land.
Habitat. — On trees or more rarely rocks in open
forest up to 2000 m elevation.
Remarks. — This species represents the isidiate
morph of P. zambiensis (Hale) Hale. It also repre-
sents the usnic acid-containing counterpart of P.
concrescens, a species which does not occur outside
of Africa. It has no relationship to P. caperata, as
the name might be construed to imply, which con-
tains protocetraric acid.
Specimens Examined. — Ivory Coast: Seguela, Santesson
10698a, 10702b (UPS, US). Guinea: N’Zerekore, Santesson
10568a (UPS, US). Zaire: Hoeg (TRH), Schmitt 3048 (BR).
Angola: Bie, Degelius (Degelius herbarium); Cuanza Sul,
Degelius (Degelius herbarium); Huila, Degelius (Degelius
herbarium); Mexico, Degelius (Degelius herbarium). Rho-
desia: Angus (BM), Hoeg (TRH), Kofier (LD), Schiitte 61b
(LD, US). Malawi: Jellicoe 55 (BM). Nepal: Poelt 107, 114
(M, US), Noordyte (L), Togashi (TNS). India: Tamil Nadu,
Awasthi 4431, 4432 (Awasthi herbarium, US), Degelius As-
256 (Degelius herbarium), Foreau 54 (Awasthi herbarium,
US), Hale 43673, 43713, 43866, 43948; Uttar Pradesh, Awasthi
3496, 3972 (Awasthi herbarium). Thailand: Kurokawa 1670,
1954 (TNS, US), Tsuyama (TNS, US).
Pseudoparmelia epilenca
Figure 11a
Pseudoparmelia epileuca (Hale) Hale, 1974:190.
Parmelia epileuca Hale, 1972:343 [type collection: District
Inhambane, Mozambique, Schelpe 4461 (BOL, holotype;
isotypes in LD, US)].
Descriptions. — Thallus closely adnate on bark,
ashy white, 3-5 cm broad; lobes sublinear, contigu-
ous, with smooth dark rimmed margins, 1. 5-2.0 mm
wide; upper surface plane, sorediate, the soralia
orbicular, confluent with age; lower surface black,
moderately rhizinate. Apothecia unknown.
Chemistry. — Cortex K+ yellow, medulla K — ,
C — , KC — , P+ red; atranorin and protocetraric
acid.
Distribution. — Kenya and Mozambique.
Habitat. — On coconut and other trees in open
forest or savanna up to 300 m elevation.
Remarks. — This rare lichen appears to be the
sorediate morph of P. schelpei (Hale) Hale, which
also occurs in Mozambique. Two superficially re-
lated sorediate species with protocetraric acid occur
in the New World; P. alabamensis (Hale and Mc-
Cullough) Hale, which is saxicolous and has gen-
erally narrower lobes, and P. raukiaeri (Vainio)
Hale, which has irregular pustular soredia.
Specimens Examined. — Kenya: Coast Province, Santesson
20898 (UPS, US), Mozambique: Sul do Save, Schelpe 4461a
(BOL, US), 4460a, 4461b (BOL).
Pseudoparmelia eruptens
Figure 116
Pseudoparmelia eruptens (Kurokawa) Hale, 1974:190.
Parmelia eruptens Kurokawa in Hale and Kurokawa, 1964:
153 [type collection: Lydenburg, Transvaal, Union of South
Africa, Almborn 7498 (LD, holotype; US, isotype)].
Description. — Thallus adnate, corticolous, whit-
ish mineral gray to buff in the herbarium, 5-8 cm
broad; lobes subirregular, rotund, 2-8 mm wide;
upper surface plane, continuous, moderately isidiate-
pustulate, the isidia irregularly inflated, basally
constricted, bursting apically; lower surface black,
sparsely rhizinate except for a narrow naked zone
at the tips. Apothecia rare, adnate, 1-3 mm in
diameter, the amphithecium coarsely isidiate; spores
8, 5-7 X 19-12 /cm.
Chemistry. — Cortex K+ yellow, medulla K — ,
C — , KC— or KC+ purple violet, P — ; atranorin
and divaricatic acid with associated unknowns.
Distribution. — Mozambique and Union of South
Africa.
Habitat. — On trees (and rocks?) in open forest.
Remarks. — This rare species is probably most
closely related to sorediate P. texana (Tuckerman)
Hale. The large pustules of P. eruptens do not be-
come sorediate.
Specimens Examined. — Mozambique: Mitchell 332 (US).
Union pf South Africa: Transvaal, Maas Geesteranus 6453
(L, US), 6455 (L).
30
SMITHSONIAN CONTRIBUTIONS TO BOTANY
lllllllllllllllillllllSM
Figure 11. — Species of Pseudoparmelia: a, P. epileuca ( Schelpe 4461b, isotype in BOL); b, P.
eruptens (Almborn 7498, holotype in LD); c, P. exornata ( Lamb 1101 in US); d, P. ferax ( Rogers
1326 in US); e, P. geesterani (Maas Geesteranus 6405, holotype in L); f, P. gerlachei (Santesson
6495 in US). (Scale in mm.)
NUMBER 31
31
Pseudoparmelia exornata, new combination
Figure 11c
Parmelia caperata var. exornata Zahlbruckner, 1912:379 [type
collection: Cerillos Canelones, Uruguay, Felippone 431 (W,
lectotype; G, isolectotype)].
Parmelia rutidota f. filizans Lynge, 1914:153 [type collection:
Quinta, Rio Grande do Sul, Brazil, Malme 727 (S, lecto-
type)].
Description. — Thallus adnate to closely adnate
on bark, yellowish green, 4-10 cm broad; lobes sub-
irregular, contiguous, apically subrotund, 2-4 mm
wide, sometimes marginally dissected, becoming
lobulate and crowded toward the center, with nu-
merous pycnidia; lower surface black and sparsely
rhizinate but with a rugose bare brown zone at the
margin near the tips. Apothecia common, adnate
to substipitate, 2-5 mm in diameter, the rim cre-
nate; spores 8, 8-10 X 15-18 p.m.
Chemistry. — Cortex K — , medulla K — , C — ,
KC— or KC+ rose, P+ orange; usnic acid, proto-
cetraric acid, and the “conformata” unknown.
Distribution. — Southeastern Brazil and Uruguay.
Habitat. — On trees in open forest at 300-600 m
elevation.
Remarks. — This species is closely related to P.
rutidota in chemistry and morphology. It produces
an unidentified substance, the “conformata” un-
known, just below protocetraric acid in the hexane
solvent, the same chemistry known for its presump-
tive isidiate morph, the saxicolous P. papillosa. The
lobes tend to be thinner and more filiform and
finely lacinate than typical P. rutidota. It occupies
a restricted range in Brazil and Uruguay, whereas
P. rutidota occurs sporadically in the Andean chain
and in northern Brazil, as well as in North America
and Australia.
Specimens Examined. — Brazil: Rio Grande do Sul, Malme
715 (S). Uruguay: Lavalleja, Lamb 1101 (FH, US); San Jose,
Osorio 877, 6154, 6195 (MVM); Tacuarembo, Osorio 1106,
1165 (MVM); Treinta y tres, Osorio 5979 (MVM).
Pseudoparmelia ferax
Figure lid
Pseudoparmelia ferax (Muller Argoviensis) Hale, 1974:190.
Parmelia ferax Muller Argoviensis, 1886:257 [type collection:
Australia: New South Wales, Gutawang, Hamilton 2 (G,
lectotype)].
Parmelia obversa Stirton, 1899:76 [type collection: Australia,
Paton (GLAM, lectotype; BM, isolectotype)].
Parmelia citrinescens Gyelnik, 1938b:271 [type collection:
Lago Nahuel, Puerto Blest, Patagonia, Argentina, Dusen
163 (BP, holotype; S, US, isotypes)].
Description. — Thallus closely adnate on bark,
greenish yellow, 5-8 cm broad; lobes subirregular,
crowded, apically rotund, 3-4 mm wide; upper sur-
face soon wrinkled and rugose, in part warty and
lobulate; lower surface black and coarsely rhizinate
except for a narrow brown zone at the margins.
Apothecia common, sessile, plane to almost urceo-
late, 2-4 mm in diameter; spores 8, 7-8 X 13-16 ^m.
Chemistry. — Cortex K+ yellowish, medulla K — ,
C — , KC — , P+ red; atranorin, usnic acid, and
physodalic acid.
Distribution. — Australia and Chile.
Habitat. — On branches and trunks of shrubs and
trees in arid habitats up to 1200 m elevation.
Remarks. — This species was almost always iden-
tified as “Parmelia rutidota” until the chemistry
was clarified by Kurokawa (1967). Pseudoparmelia
rutidota contains protocetraric acid and generally
has a more expanded thallus. While described from
Australia, P. ferax seems to be most common in
Chile.
Specimens Examined. — Australia: South Australia, Rogers
1320, 1326 (US); Victoria, Filson 6597 (US). Chile: Aconcagua,
Follmann 11784-L (US); Nuble, Mahu 3538 (US); Santiago,
Mahu 1117, 2034, 3334 (US), Santesson 7124 (S, US); Valpa-
raiso, Imshaug 36657 (MSC), Rundel 7335 (US).
Pseudoparmelia geesterani
Figure lie
Pseudoparmelia geesterani (Hale) Hale, 1974:190.
Parmelia geesterani Hale, 1972b:344 [type collection: Trans-
vaal, Union of South Africa, Maas Geesteranus 6405 (L,
holotype; LD, US, isotypes)].
Description. — Thallus closely adnate on rock,
dark olive greenish to whitish, 2-4 cm broad; lobes
sublinear, crowded, 0. 5-1.0 mm wide; upper surface
plane, isidiate, isidia coarse and irregularly inflated;
lower surface brown or blackening, sparsely rhizi-
nate. Apothecia unknown.
Chemistry. — Cortex K+ yellow, medulla K +
yellow turning red, C — , KC — , P+ orange; atrano-
rin and salazinic acid.
Distribution. — Union of South Africa.
Habitat. — On exposed rocks at 1500-1800 m ele-
vation.
Remarks. — This lichen is still known only from
32
SMITHSONIAN CONTRIBUTIONS TO BOTANY
the type collection. It is not closely related to any
other Pseudoparmeliae although it might be con-
fused with isidiate-pustulate P. owariensis (Asahina)
Hale or P. imperfecta (Kurokawa) Hale.
Pseudoparmelia gerlachei
Figure 11/
Parmelia gerlachei Zahlbruckner, 1929:137 [type collection:
based on Parmelia antarctica Vainio].
Parmelia antarctica Vainio, 1903:13 [type collection: Cap van
Beneden, Terre de Danco, Gerlache 226 (TUR, Vainio
herbarium number 2839, lectotype); not P. antarctica Bit-
ter, 1901:248 (= Hypogymniq)].
Xanthoparmelia gerlachei (Zahlbruckner) Hale, 1974:487.
Description. — Thallus closely adnate on rock,
rather coriaceous, pale greenish yellow, 3-5 cm
broad; lobes subirregular, crowded, apically rotund,
2-4 mm wide; upper surface dull, sometimes lightly
pruinose, broadly rugose or foveolate, appearing
somewhat inflated, soredia developing in orbicular
to irregular laminal, capitate soralia 2-4 mm in
diameter; lower surface black and moderately
rhizinate, papillate in a narrow brown marginal
zone. Apothecia not found.
Chemistry. — Cortex K — , medulla K — , C — ,
KC — , P+ red; usnic acid and either physodalic
acid or protocetraric acid or a mixture of both.
Distribution. — Andean mountain chain and
Antarctica.
Habitat. — On rocks in exposed paramo or tundra
at 2900-3800 m in the Andes and to sea level in
Antarctica.
Remarks. — I had previously considered this to be
a Xanthoparmelia because of the saxicolous habitat
and presence of usnic acid. The apically rotund lobe
configuration and penicillate rhizines and papillae
at the margin, as well as physodalic acid, are more
characteristic of Pseudoparmelia, as suggested by
Kurokawa (1967). Pseudoparmelia gerlachei has
evolved in extremely harsh environments where
trees are completely absent. The most closely related
species and the probable progenitor is P. ferax
(Muller Argoviensis) Hale, which occurs on trees in
Australia and Chile. Both species contain physodalic
acid in the “typical” state, but P. gerlachei in Ven-
ezuela contains only protocetraric acid whereas
Chilean and Argentinian specimens produce both
acids. This combination is similar to that in Hypo-
trachyna physodalica (Hale) Hale (Hale, 1975a).
Specimens Examined. — Venezuela: Merida, Hale 42706,
42843, 44667, 44676. Chile: Magallanes, Santesson 1924 (S, US).
Argentina: Santa Cruz, Santesson 7079 (S, US). Antarctica:
Graham Land, Lamb 2644 (FH, US); Cap van Beneden, Ger-
lache (TUR, Vainio herbarium numbers 2840, 2841, syntypes
of P. antarctica).
Pseudoparmelia hypomilta
Figure 12 a
Pseudoparmelia hypomilta (Fee) Hale, 1974:190.
Parmelia hypomilta Fee, 1837:123 [type collection: Peru (G,
lectotype; H, isolectotype)].
Parmelia regnellii Lynge, 1914:40 [type collection: Sao Joao
d’el Rey, Minas Gerais, Brazil, Malme 308 (S, lectotype;
BM, MICH, isolectotypes)].
Parmelia regnellii f. arida Lynge, 1914:141 [type collection:
Bocca da Serra, Mato Grosso, Brazil, Malme 2240 (S, lecto-
type; UPS, US, isolectotypes)].
Description. — Thallus closely adnate on bark,
rather coriaceous, buff to straw colored, 1-3 cm
broad; lobes sublinear 1.0-1. 5 mm wide, often black
rimmed; upper surface plane to convex, densely
white maculate; medulla white in upper half, dull
red in the lower half; lower surface brown, moder-
ately rhizinate, the rhizines brown. Apothecia (de-
scription from Malme 171) common, adnate, 1-2
mm in diameter; spores 8, more or less uniseriate,
5X6 fim.
Chemistry. — Cortex K+ yellowish, medulla K + ,
C T , KC+ yellowish (pigment K-f purple), P — ;
atranorin and unidentified substances.
Distribution. — Brazil.
Habitat. — On trees and rocks in open forests.
Remarks. — Pseudoparmelia hypomilta is a vari-
able species in terms of lobe width. Lynge’s f. arida
has quite narrow lobes (1 mm or less) but the types
of P. hypomilta and Parmelia regnellii are com-
parable in size. While it is difficult to generalize on
a species so poorly represented in herbaria, it seems
closely related to P. congruens but is readily differ-
entiated by the medullary pigment and, as far as we
can determine, common occurrence on rock.
Specimens Examined. — Brazil: Minas Gerais, Gardner
(BM), Malme 171 (LD, UC, US), Warming 302 (M); Mato
Grosso, Malme 2240 (S,US), Malme in Lichenes Austroameri-
cani 92 (G, LD, S, UPS).
Pseudoparmelia inhaminensis
Figure 12b
Pseudoparmelia inhaminensis (Dodge) Hale, 1974:190.
Parmelia inhaminensis Dodge, 1959:130 [type collection: In-
hambane, Inhamine, Angola, Sousa (BM, holotype)].
NUMBER 31
33
rv
,Tw#'f
Figure 12. — Species of Pseudoparmelia: a, P. hypomilta (Malme 2745B in US); b. P. inhamin-
ensis (Degelius in US); c, P. inornata (Imshaug 24454, isotype in US); d, P. intertexta ( Pentype ,
lectotype of P. gracilis Muller Argoviensis in G); e, P. -ischnoides (Almborn 1698, holotype in
LD); /, P. labrosa (James 577 in US). (Scale in mm.)
34
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Description. — Thallus adnate on twigs, soft and
fragile, buff mineral gray, 3-5 cm broad; lobes sub-
irregular, apically rotund, 1.5-2 mm wide; upper
surface more or less regularly rugose and wrinkled,
the cortex easily breaking away, heavily pycnidiate;
lower surface black and moderately rhizinate except
for a narrow brown, bare zone at the tips. Apothecia
substipitate, 2 mm in diameter; spores 8, 5-8 X
10-12 /xm.
Chemistry.— Cortex K -1- yellow, medulla K +
yellow, C — , KC — , P-f orange; atranorin, stictic
acid, and constictic acid.
Remarks. — The chemical constituents and rugose
upper surface place P. inhaminensis close to P. cro-
zalsiana (Bouly de Lesdain) Hale, which is sorediate,
and P. scrobicularis (Krempelhuber) Hale, which
is nonsorediate and has a pruinose apothecial disc,
a smaller, very rugose thallus, and unusually large
conidiospores (about 20 /xm long). The conidio-
spores of P. inhaminensis are about 12/xm long. It is
a possible candidate as nonsorediate progenitor of
P. crozalsiana.
Specimens Examined. — Angola: Mo^araedes, Degelius (De-
gelius herbarium, US).
Pseudoparmelia inornata
Figure 12c
Pseudoparmelia inornata (Hale) Hale, 1974:190.
Parmelia inornata Hale, 1971a:32 [type collection: Grand
Cayman, Imshaug 24454 (MSC, holotype; US, isotype)].
Description. — Thallus adnate on bark, pale
greenish mineral gray, 5-10 cm broad; lobes sub-
irregular, apically rotund, 3-7 mm wide; upper
surface plane, continuous to cracked with age, usu-
ally densely pycnidiate, white-reticulate at the lobe
tips; lower surface black and sparsely rhizinate ex-
cept for a narrow bare, brown zone at the margins.
Apothecia common, substipitate, 2-4 mm in diam-
eter; spores 8, 7-8 X 16—18 /xm.
Chemistry. — Cortex K+ yellow, medulla K — ,
C — , KC+ rose, P+ red; atranorin and proto-
cetraric acid.
Distribution. — West Indies.
Habitat. — On trees in shaded woods near sea
level.
Remarks. — This rather broad-lobed species might
be considered as a possible progenitor of isidiate P.
martinicana (Nylander) Hale, but it is more coriac-
eous and has a different aspect. It appears to be best
developed in dry sea-level forests of smaller islands.
Specimens Examined. — Records from the Bahamas, Grand
Cayman, and Haiti are listed in Hale (1971:32).
Pseudoparmelia intertexta
Figure 12d
Pseudoparmelia intertexta (Montagne and van den Bosch)
Hale, 1974:190.
Parmelia intertexta Montagne and van den Bosch in Mon-
tagne, 1856:327 [type collection: Java, Junghuhn (L, lecto-
type; PC, isolectotype)].
Parmelia ecoronata Nylander, 1873:64 [type collection: Pulo-
Penang, Malaya, Collingham [Cunningham in publication]
(H, Nylander herbarium number 32999, lectotype)].
Parmelia subrupta Nylander in Nylander and Crombie, 1883:
51 [type collection: Allagajah, near Malacca, Malaya,
Maingay (BM, lectotype; H, isolectotype)].
Parmelia gracilis Muller Argoviensis, 1887:317 [type collec-
tion: Daintree River, Australia, Pentzke (G, lectotype); not
Parmelia gracilis (Persoon) Sprengel, 1827:277 (= Usnea)].
Parmelia relicina var. ecoronata (Nylander) Muller Argovi-
ensis, 1891:378.
Parmelia gracilenta Vainio, 1900:6 [type collection: Based on
Parmelia gracilis Muller Argoviensis].
Description. — -Thallus closely adnate, corticolous,
marguerite yellow, 3-10 cm in diameter; lobes sub-
linear-elongate, 0.5-2 mm wide; upper surface more
or less convex, faintly maculate; lower surface pale
brown to tan, densely rhizinate, the rhizines simple
to densely branched, pale. Apothecia numerous,
adnate, 0.7-2 mm in diameter; spores 8, 3-5 X 5-7
/xm.
Chemistry. — Cortex K+ yellowish, medulla K — ,
C — , KC+ rose, P+ orange red; atranorin, proto-
cetraric acid, protolichesterinic acid, and usnic acid.
Distribution. — Andaman Islands, Thailand, Ma-
laysia, Philippines, Indonesia, New Guinea, and
Australia.
Habitat. — On canopy branches of trees (diptero-
carps and Quercus) in rain forest at 150-1600 m
elevation.
Remarks. — This is one of two Pseudoparmeliae
(the other being P. malaccensis (Nylander) Hale)
that have evolved in the Southeast Asian rain for-
ests, primarily on dipterocarps. The only other com-
mon parmelioid genus there is Relicina (Hale,
1975b), which is also characterized by a closely
adnate habit and presence of usnic acid. Pseudo-
parmelia intertexta is a presumptive nonisidiate
progenitor of P. malaccensis (Nylander) Hale (see
NUMBER 31
35
discussion under that species), although it is anom-
alous in having branched rhizines.
Specimens Examined. — Andaman Islands: Kurz 3 (M, UPS,
W). Thailand: Chang, Schmidt XVI (TUR). Malaya: Pahang,
Hale 30184, 30225, 30482, 30498; Selangor, Hale 30073, 30260,
30261, 30264, 30299, 31188. Philippines: Mountain Province,
Hale 25811; Negros Occidental Province, Hale 26422, 26511,
26523. Malaysia: Sabah, Hale 28206, 28813, 29065, 30363,
30364. New Guinea: Versteegh (L, US).
Psendoparmelia ischnoides
Figure 12a
Pseudoparmelia ischnoides (Kurokawa) Hale, 1974:190.
Parmelia ischnoides Kurokawa in Hale and Kurokawa, 1964:
155 [type collection: Window Stream, Kirstenbosch, Wyn-
berg. Cape Province, Union of South Africa, Almborn
1698 (LD, holotype; TNS, US, isotypes)].
Description. — Thallus closely adnate on rock,
fragile, whitish ashy gray, 4-10 cm in diameter;
lobes sublinear-elongate, 0.5-2 mm wide; upper sur-
face plane to convex, continuous, isidiate, the isidia
short, simple, darkening at the tips; lower surface
black, sparsely rhizinate. Apothecia adnate, 1-2 mm
in diameter, the amphithecium isidiate; spores 8,
5-6 X 7-8 p.m.
Chemistry. — Cortex K+ yellow, medulla K +
yellow, C — , KC — , P+ pale orange; atranorin, stic-
tic acid, and constictic acid.
Distribution. — Union of South Africa.
Habitat. — On rocks in open areas.
Remarks. — This species has a very limited range
in South Africa and probably represents a typical
Cape endemic. It bears a superficial resemblance to
two other small saxicolous species, P. annexa (Kuro-
kawa) Hale (lecanoric acid present) and P. arcana
(Kurokawa) Hale (fatty acids, pale below).
Specimens Examined. — See Hale and Kurokawa (1964:156)
for records in the Union of South Africa.
Pseudoparmelia labrosa
Figure 12/
Pseudoparmelia labrosa (Zahlbruckner) Hale, 1974:190.
Parmelia tenuirima var. labrosa Zahlbruckner, 1941:108 [type
collection: Saddle Hill, Dunedin, New Zealand, Thomson
V34 (W, lectotype)].
Parmelia labrosa (Zahlbruckner) Hale, 1968:325.
Description. — Thallus adnate on bark, light buff
mineral gray, 4-8 cm broad; lobes subirregular.
apically subrotund, 1.5-3 mm wide; upper surface
plane to rugulose, shiny, sorediate, the soralia orig-
inating from coarse pustular ridges, becoming irregu-
lar to diffuse; lower surface black and moderately
rhizinate. Apothecia adnate, 3-5 mm in diameter;
the rim sorediate; spores 8, 5 X 10-12 p,m.
Chemistry. — Cortex K+ yellow, medulla K — ,
C T, KC+ red, P — ; atranorin and lecanoric acid.
Distribution. — Australia, New Zealand, and
Chile.
Habitat. — On trees ( Drachophyllum , Myrsine,
Hymenanthera, and Betula) in open woods at low
elevations.
Remarks. — The lobe configuration places this
austral species near P. texana (Tuckerman) Hale,
but the soralia are more diffuse and the chemistry is
distinct. The parallel isidiate morph is probably
P. conlabrosa Hale; there seems to be no fertile
progenitor. The syntype of P. tenuirima var. labrosa
( Thomson 2A 683 in W) is Parmotrema reticulatum
(Taylor) Choisy.
Specimens Examined. — Chile: Chiloe, Santesson 2262 (S).
Other records from Australia and New Zealand are listed in
Hale (1968:325).
Pseudoparmelia lecanoracea
Figure 13a
Pseudoparmelia lecanoracea (Muller Argoviensis) Hale, 1974:
190.
Parmelia lecanoracea Muller Argoviensis, 1888:529 [type col-
lection: Arisdrift, Oranje River, Namaqualand, Union of
South Africa, Schenck 543 (G, lectotype)].
Description. — Thallus closely adnate, appearing
areolate at the center, pruinose whitish buff, 2-3 cm
broad; lobes sublinear, 0. 6-1.0 mm wide, black
rimmed; upper surface convex, rugose with age,
roughened; medulla pigmented reddish yellow in
the lower half; lower surface tan or darkening, mod-
erately rhizinate, the rhizines brown. Apothecia
rare, adnate, 1 mm in diameter; spores 8, 7 X 8-10
/xm.
Chemistry. — Cortex K+ yellow, medulla K — ,
CT, KC+ rose; atranorin and evernic acid; pig-
mented medulla K+ purple, skyrin present.
Distribution. — Union of South Africa.
Habitat. — On rocks.
Remarks. — This species is still only known from
the rather fragmentary type collection. As Muller
noted, it could be confused with a Lecanora. The
36
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Figure 13. — Species of Pseudoparmelia: a, P. lecanoracea ( Schenck 543, lectotype in G); b,
P. leucoxantha ( Eiten 4497A in US); c, P. malaccensis (Hale 24859); d, P. martinicana ( Hale
35662); e, P. molybdiza (Degelius SA-39 in US); /, P. nairobiensis (Swinscow 6/1970 n US).
(Scale in mm.)
NUMBER 31
37
chemistry is anomalous for the genus. It is included
here in the hope that future workers will be in a
better position to assess its exact relationship.
Pseudoparmelia leucoxantha
Figure 13 b
Pseudoparmelia leucoxantha (Muller Argoviensis) Hale, 1974:
190.
Parmelia leucoxantha Muller Argoviensis, 1881:85 [type
collection: Brazil, Sao Paulo, Puiggari 1050 (G, lectotype;
W, isolectotype)].
Description. — Thallus adnate to loosely attached
on rock or bark, dull greenish yellow, 3-6 cm broad;
lobes subirregular, apically rotund, 3-5 mm wide;
upper surface plane, rimose with age, sorediate
along the margins and in part on the surface,
soralia irregular in capitate or elongate masses, the
soredia coarse; lower surface black and sparsely
rhizinate except for a brown, bare or papillate zone
along the margins. Apothecia rare, sessile, up to 2
mm in diameter, the amphithecium sorediate;
spores not developed.
Chemistry. — Cortex K+ yellowish, medulla K — ,
C — , KC— or KG+ rose, P+ red; atranorin (trace),
usnic acid, and protocetraric acid.
Distribution. — Mexico and Brazil.
Habitat. — On sandstone boulders, more rarely
on trees, in dry scrubby areas (chapada vegetation
in Brazil) at 300-1100 m elevation.
Remarks. — There is considerable resemblance to
Pseudoparmelia caperata (L.) Hale in lobe config-
uration, but the soralia are more discrete and dis-
tinctly marginal and lateral. It may well have
evolved from nonsorediate P. rutidota (Hooker
and Taylor) Hale but differs in substratum.
Specimens Examined. — Mexico: Oaxaca, Hale 20643, 20648.
Brazil: Maranhao, Eiten and Eiten 4497A (US); Mato Grosso,
Malme (S); Sao Paulo, Schindler 4564b (KR, US).
Parmelia malaccensis
Figure 13c
Pseudoparmelia malaccensis (Nylander) Hale, 1974:190.
Parmelia malaccensis Nylander in Nylander and Crombie,
1883:52 [type collection: St. Johns Hill, Malacca, Malaya,
Maingay 21 (BM, lectotype; FH, H, Nylander herbarium
number 34984, isolectotypes)].
Description. — Thallus closely adnate on bark.
yellowish green, 2-8 cm in diameter; lobes short,
sublinear, 0.5-1. 5 mm wide; upper surface plane
to convex, shiny, distinctly white reticulate at the
lobe tips, tangentially cracked with age, isidiate, the
isidia simple, 0.2-0. 5 mm high; lower surface pale
brown, densely rhizinate, the rhizines simple to
sparsely branched, becoming dark brown. Apo-
thecia rare, adnate, 1-2 mm in diameter; spores 8,
4-5 X 6-7 /j,m.
Chemistry. — Cortex K+ yellowish, medulla K — ,
C — , KC+ rose, P-f red; protocetraric acid and
usnic acid.
Distribution. — Africa, India, Sri Lanka, Indo-
nesia, Malaysia, and the Philippines.
Habitat. — On trunks and canopy branches of
trees in lowland rain forest from sea level to 150 m
elevation.
Remarks. — While I tentatively consider this as
the isidiate morph of P. intertexta, mostly because
of the chemical characters and similar habitat in the
lowland rain forest of Southeast Asia, the two
species have diverged considerably in rhizine struc-
ture. The rhizines of P. malaccensis are simple and
often turn darker brown that the lower cortex; they
are pale and rather richly branched in P. intertexta.
The upper cortex is strongly white-reticulate in P.
malaccensis, continuous in P. intertexta.
Specimens Examined. — Ivory Coast: Santesson 10397a (UPS,
US). India: Tamil Nadu, Hoeg (Awasthi herbarium, US).
Sri Lanka: Fosberg 51032 (US). Philippines: Basilan, Hale
24941, 25331; Zamboanga del Norte, Hale 24727; Zamboanga
del Sur, Hale 24859. Malaya: Selangor, Hale 30055, 30068,
30262, 30266, 30284, 30292, 30295, 31187. Sarawak: Hale 29992,
29994, 29998, 30000. Sabah: Hale 30353, 30362, 30368. Indo-
nesia: Java, Spanjaard 6102 (L).
Parmelia martinicana
Figure 13d
Pseudoparmelia martinicana (Nylander) Hale, 1974:190.
Parmelia martinicana Nylander, 1885:609 [type collection:
Martinique, Tar din (H, lectotype)].
Description. — Thallus adnate, 3-9 cm broad,
pale tan mineral gray; lobes irregular, subrotund,
short, 2-5 mm wide; upper surface becoming rugose
toward the center, densely isidiate, isidia initially
papillate, cylindrical to irregularly thickened, sim-
ple or branched, rarely turning granular at the tips;
lower surface black and sparsely rhizinate at the
center, rugose, brown, and naked in a narrow zone
at the margin. Apothecia not seen.
38
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Chemistry. — Cortex K+ yellowish, medulla K — ,
C + , KC+ rose, P+ orange red; atranorin, gyro-
phoric acid, and protocetraric acid (rarely with
norlobaridone).
Distribution. — Southeastern United States, Mex-
ico, West Indies, and Venezuela.
Habitat. — On tree trunks and branches in pas-
tures and secondary forests from sea level to 1000
m elevation.
Remarks. — This is one on the commonest foliose
lichens in scrub forest and dry savanna in the West
Indies. It may be related to nonisidiate P. inornata
(Hale) Hale but has a thinner, more fragile thal-
lus. It is most closely related to P. raunkiaeri
(Vainio) Hale, which is distinctly coarsely pus-
tulate-sorediate. One specimen (Allard 17325) con-
tained norlobaridone in addition to protocetraric
acid, but further collections should be examined
before deciding whether this population represents
a distinct species or not.
Specimens Examined. — United States: Florida, Hale 36813,
36864. Mexico: Veracruz, Hale 19743. Bahamas: Merrill (US),
Britton 942, 1080 (FH). Cuba: Pinar del Rio, Imshaug 25289
(MSC); Guantanamo, Hioram 2701 (US); Oriente, Hioram
5477, 5706 (US). Dominican Republic: Allard 17325 (US),
Wetmore 3922 (MSC), 3947 (MSC, US). Puerto Rico: Britton
1691 (FH, NY). St. Croix: Britton 75 (NY, US), Boergessen
(C), Raunkiaer 547 (C), Paulsen (C). St. Martin: Le Gallo
478b, 571 (US). St. Barthelemy: Le Gallo 415, 497, 515, 521,
542a, 545, 563 (US). Guadeloupe: Culberson 14536, 14544
(DUKE), Degelius (Degelius herbarium) Duss (C), 489 (NY),
Le Gallo 465, 471, 489, 574, 575, 580 (US), 2729 (MSC), 2741
(BM). Dominica: Imshaug 33159, 33282 (MSC) (other records
listed in Hale, 1971:18). Martinique: Culberson 14732 (DUKE),
Degelius (Degelius herbarium), Jardin (H). St. Lucia: Imshaug
29701, 29968, 30144 (MSC), Evans 75 (FH, NY, YU), 101
(US). St. Vincent: Elliott (TUR), Imshaug 30369, 30692
(MSC). Tobago: Imshaug 31565 (MSC). Trinidad: Broadway
8099 (BM). Venezuela: Magdalena, Broadway 805 (US).
Pseudoparmelia molybdiza
Figure 13e
Pseudoparmelia molybdiza (Nylander) Hale, 1974:190.
Parmelia molybdiza Nylander in Crombie, 1876a: 19 [type
collection: Table Mountain, Cape of Good Hope, Union of
South Africa, Eaton (BM, lectotype; H, Nylander herbarium
number 35234, isolectotype)].
Parmelia atrichoides Nylander in Crombie, 1876b: 167 [type
collection: Cape of Good Hope, Union of South Africa,
Eaton (BM, lectotype; H, isolectotype)].
Parmelia brachyphylla Muller Argoviensis, 1886:256 [type
collection: Near Lydenburg, Transvaal, Union of South
Africa, Wilms 2752 (BM, lectotype).
Parmelia perfissa Steiner and Zahlbruckner, 1926:519 [type
collection: Port Elizabeth, Cape Province, Union of South
Africa, Brunnthaler (W, lectotype; WU, isolectotype)].
Description. — Thallus adnate to appressed on
rock, whitish mineral gray, 4-8 cm broad; lobes sub-
irregular, apically rotund, crowded toward the
thallus center, 2-3 mm wide; upper surface plane,
rimose with age; lower surface dark brown and
blackening, moderately rhizinate. Apothecia numer-
ous, adnate, 1-2.5 mm in diameter; spores 8, 5 X 6
/xm.
Chemistry. — Cortex K+ yellow, medulla K — ,
C T , KC + red, P — ; atranorin and lecanoric acid.
Distribution. — Uganda and Union of South
Africa.
Habitat.- — On rocks in open or partly shaded
habitats from sea level to about 1200 m elevation.
Remarks. — Pseudoparmelia molybdiza is espe-
cially common in Cape Province. The isidiate morph
appears to be P. annexa (Kurokawa) Hale. The
brilliant C -f red test identifies it immediately. Pseu-
doparmelia spodochroa (Kurokawa and Filson)
Hale, P. tortula (Kurokawa) Hale, and even P.
vanderbylii (Zahlbruckner) Hale are very similar
externally and occur in the same localities where
P. molybdiza is found. They would be differen-
tiated by a negative C test.
Specimens Examined. — Uganda: Pian County, Swinscow
2U 31/10A (BM, US). Union of South Africa: Natal, Aim-
born 8645, Hoeg (TRH); Transvaal, Maas Geesteranus (L);
Basutoland, Kofler (LD); Orange Free State, Almborn 5818,
5831, 5834 (LD); Cape Province, Almborn 1803, 2065, 2066,
2067 , 4837, 4948, 4980, 5690, 11115 (LD), Degelius SA-39
(US), Hoeg (LD, TRH), Maas Geesteranus 6706, 6707, 6730
(L).
Pseudoparmelia nairobiensis
Figure 13/
Pseudoparmelia nairobiensis (Steiner and Zahlbruckner) Hale,
1974:190.
Parmelia nairobiensis (“neirobiensis’”) Steiner and Zahlbruck-
ner, 1926:517 [type collection: Nairobi, Kenya, Schroder
287 (W, lectotype)].
Parmelia gracilescens var. angolensis Vainio in Welwitsch,
1901:401 [type collection: Serra da Xella, Huila, Angola,
Welwitsch 30 pro parte (TUR, Vainio herbarium number
3059, lectotype)].
Parmelia angolensis (Vainio) Dodge, 1959:103.
Parmelia ganguellensis Dodge, 1959:109 [type collection:
Ganguelas and Ambuelas, Benguela, Angola, Gossweiler
(BM, lectotype)].
NUMBER 31
39
Parmelia hansfordii Dodge, 1959:127 [type collection: Kam-
pala, Uganda, Hansford 1455 (BM, lectotype)].
Description. — Thallus loosely adnate to ap-
pressed on bark, rather coriaceous, greenish to buff
mineral gray, 5-10 cm broad; lobes sublinear to sub-
irregular, usually apically subrotund, 2-5 mm wide,
often becoming marginally lobulate with age; upper
surface plane, shiny, usually conspicuously pycni-
diate, reticulately rimose with age; lower surface
black except for a narrow marginal brown zone,
sparsely to moderately rhizinate. Apothecia com-
mon, adnate, 2-5 mm in diameter; spores 8, 6-8 X
8-13 fxm.
Chemistry. — Cortex K+ yellow, medulla K — ,
C — , KC+ faint violet or KC — , P — ; atranorin and
divaricatic acid with associated unknowns.
Distribution. — Kenya, Uganda, Angola, Zaire,
Rhodesia, and Tanzania.
Habitat. — On trees and rocks in open or second-
ary forest at 1000-1700 m elevation.
Remarks. — This is one of the more common foli-
ose lichens in central Africa. It appears to be (or
at least would be similar to) the progenitor for
isidiate P. concrescens (Vainio) Hale and sorediate
P. texana (Tuckerman) Hale.
Specimens Examined. — Uganda: Buddu County, Lye L167
(BM, US); Burahiya County, Swinscow 2U 17/21 (BM, US).
Kenya: Central Province, Maas Geesteranus 10272 in Lichenes
Africani 29 (L, LD, US). Zaire: Degelius (Degelius herbar-
ium), Louis 4857E, 4858 (BR). Tanzania: Kilimanjaro Prov.,
Santesson 20959 (UPS). Angola: Bie, Degelius (Degelius her-
barium); Huila, Degelius (Degelius herbarium); Moxico,
Degelius (Degelius herbarium). Rhodesia: Bankart (BM),
Bullock 2105 pars, Hoeg (TRH). Tanzania: Proctor 1023 B
(BM).
Pseudoparmelia neoquintaria, new species
Figure 14a
Description. — Thallus laxe adnatus, saxicola,
cinereo-albidus, 3-7 cm latus, lobis sublinearibus,
1-2 mm latis, margine laciniatis; superne planus,
continuus vel aetate rimosus, isidiatus, isidiis cylin-
dricis, simplicibus vel ramosis, usque ad 0.5 mm
altis; cortex superior 10-15 p. m crassus, epicorticatus,
epicortice perforato, stratum gonidiale 10-15 pjm.
crassum, medulla alba, ca 200 ju,m crassa, cortex in-
ferior paraplectenchymatus, 10-12 p.m crassus; sub-
tus castaneus, modice rhizinosus, rhizinis elongatis,
simplicibus, castaneis. Apothecia ignota.
Chemistry. — Cortex K + yellow, medulla K —
or K+ reddish, C — , KC — , P — ; atranorin and the
“quintaria” unknowns.
Holotype. — Australia: New South Wales, Mount
Wilson, Blue Mountains, G.E and G. Du Rietz 781,
12 November 1926 (UPS; US, isotype).
Distribution. — Australia.
Habitat. — On exposed sandstone rocks.
Remarks. — The thallus of this unique species is
rather loosely attached with a tendency for the lobes
to become revolute, a kind of growth form often
seen with soil-inhabiting Xanthoparmeliae growing
under harsh conditions. The “quintaria” unknowns
have been discovered in the brown Parmeliae and
in Xanthoparmelia quintaria (Hale) Hale, which
has quite different lobe configuration and a black
lower surface.
Pseudoparmelia owariensis
Figure 14 b
Pseudoparmelia owariensis (Asahina) Hale, 1974:190.
Parmelia owariensis Asahina, 1953:135 [type collection:
Inuyama, Province Owari, Japan, Asahina (TNS, lecto-
type)].
Description. — Thallus closely adnate on rocks,
whitish mineral gray, 2-5 cm broad; lobes sublin-
ear, 0.5-2 mm wide; upper surface plane, continuous
or cracked with age, isidiate-pustulate, the isidia
coarse, short, cylindrical to irregularly inflated,
bursting open apically; lower surface black, sparsely
rhizinate. Apothecia unknown.
Chemistry. — Cortex K+ yellow, medulla K — ,
C — , KC+ faint purple violet, P — , atranorin, and
divaricatic acid with associated unknowns.
Distribution. — Africa, Thailand, Hong Kong,
and Japan.
Habitat. — On rocks in open areas at lower eleva-
tions.
Remarks. — Pseudoparmelia owariensis is a mem-
ber of a saxicolous pustulate complex that also in-
cludes P. pustulescens. These two species cannot be
distinguished except by chemical tests. Many more
specimens will have to be examined to comprehend
more fully their relationships.
Specimens Examined . — Ivory Coast: Segu61a, Santesson
10713a (UPS). Uganda: Nyabushozi County, Swinscow 2U
22/2A, 2Z 22/4 (BM, US). Thailand: Kurokawa 1718, 1719, 1720,
1723, 1724, 1814, 1874 (TNS). Hong Kong: Thrower 1189 (BM).
40
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Figure 14. — Species of Pseudoparmelia: a, P. neoquintaria (Du Rietz 781, holotype in US);
b, P. owariensis (Kurokawa 1874 in US); c, P. pachydactyla (Kofler in US); d, P. papillosa
(Zorron 2334 in US); e, P. prolata (Hdeg, holotype in TRH); /, P. pustulescens (Santesson
10691b in US). (Scale in mm.)
NUMBER 31
41
Pseudoparmelia pachydactyla
Figure 14c
Pseudoparmelia pachydactyla (Hale) Hale, 1974:191.
Parmelia caperata var. isidiophora Steiner, 1897:215 [type
collection: Athi Plains, Kenya, Liechtenstein (WU, lecto-
type)].
Parmelia pachydactyla Hale, 1972b:345 [type collection:
based on P. caperata var. isidiophora Steiner].
Parmelia steineri Dodge, 1959:125 [type collection: based on
P. caperata var. isidiophora Steiner; not P. steineri Gyel-
nik, 1938b:289 (= Xanthoparmelia molliuscula (Acharius)
Hale)].
Description. — Thallus closely adnate on rock,
light greenish yellow, 4-6 cm broad; lobes sublinear,
crowded toward the center of the thallus, 1-1.5 mm
wide; upper surface plane, dull, isidiate, the isidia
scattered, thick, about 0.3 mm wide and to 0.5 mm
high; lower surface black and velvety to the margin,
sparsely rhizinate, the rhizines coarse, dull. Apo-
thecia unknown.
Chemistry. — Cortex K — , medulla K — , C — ,
KC— or KC+ rose, P+ red; usnic acid and proto-
cetraric acid.
Distribution. — Kenya and Rhodesia.
Habitat. — On rocks in semiarid regions at mid
elevation.
Remarks. — The black velvety lower surface is
identical to that of Pseudoparmelia amplexa (Stir-
ton) Hale, the presumptive parent species. The
sorediate morph is P. subamplexa Hale. Pseudopar-
melia pachydactyla is different from both of these
species in being saxicolous.
Specimens Examined. — Rhodesia: Division Victoria, Rol-
ler (LD, US).
Pseudoparmelia papillosa, new combination
Figure 14d
Parmelia papillosa Lynge ex Gyelnik, 1935:43 [type collec-
tion: Canelones, La P.az, Uruguay, F. Felippone 752 (W,
lectotype)].
Xanthoparmelia papillosa (Lynge ex Gyelnik) Hale, 1974:
488.
Description. — Thallus adnate on rock, greenish
yellow, 4-6 cm broad; lobes subirregular, apically
rotund, 2-4 mm wide; upper surface plane and
shiny, somewhat white-reticulate at the tips, becom-
ing rugose and densely isidiate toward the center,
the isidia irregularly inflated, to 0.4 mm thick and
up to 1 mm high, solid but the tips easily eroding
away, not becoming sorediate; lower surface black
and moderately rhizinate except for a bare or papil-
late zone at the tips. Apothecia common, substipi-
tate and often appearing immersed among the
isidia, 1.5-3 mm in diameter, the amphithecium and
rim densely isidiate; spores 8, 7-8 X 16-18 ^m.
Chemistry. — Cortex K — , medulla K — , C — ,
KC — , or KC+ reddish, P+ red; usnic acid, pro-
tocetraric acid, and the “conformata” unknown.
Distribution. — Brazil, Uruguay, and Argentina.
Habitat. — On acidic rocks in open areas from
sea level to 1000 m elevation.
Remarks. — While I had recently transferred this
species to Xanthoparmelia, partly because of its saxi-
colous habit, it displays features characteristic of
Pseudoparmelia : broad, apically rotund lobes, a
bare or papillate brown zone at the tips below, rhi-
zines with brown, fibrous tips, and presence of pro-
tocetraric acid which is rare in Xanthoparmelia,
especially in New World species. As so delimited,
P. papillosa is another offshoot from the P. ruti-
dota complex differentiated by the very large almost
pustular isidia. A comparable saxicolous species con-
taining protocetraric acid, without the “confor-
mata” unknown, P. baltimorensis (Gyelnik and
Foriss) Hale, has even larger more typically pustular
outgrowths.
Specimens Examined. — Brazil: Santa Catarina, Reitz and
Klein 15977, 16120 (US). Uruguay: Lavalleja, Lamb 1107
(FH, US); Maldonado, Osorio 5026 (MVM, US); Rocha,
Herter 826a (H, S, US), Zorron 2334 (US); Treinta y Tres,
Herter 2 (H). Argentina: Buenos Aires, Eyerdam 2369b (F).
Pseudoparmelia prolata
Figure 14e
Pseudoparmelia prolata (Hale) Hale, 1974:191.
Parmelia prolata Hale, 1972:344 [type collection: Cape
Province, Union of South Africa, Hdeg (TRH, holotype;
LD, US, isotypes)].
Description. — Thallus adnate to loosely attached
on rock or rarely on soil, pale olivaceous mineral
gray, 4-7 cm broad; lobes sublinear, extended, al-
most divaricate, 2-3 mm wide; upper surface plane,
shiny; lower surface pale brown, moderately rhiz-
inate, the rhizines pale brown. Apothecia not seen.
Chemistry. — Cortex K-f yellow, medulla nega-
tive with all reagents; atranorin and an unknown
spot (aliphatic compound?).
42
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Distribution. — Union of South Africa.
Habitat. — On rocks, more rarely on loose soil,
in open areas.
Remarks. — The general configuration of this
species is that of a Xanthoparmelia. In many re-
spects it resembles P. neoquintaria Hale, which
occurs in similar habitats in Australia but has a
different chemistry. The isidiate morph is P. basu-
toensis (Hale) Hale.
Specimens Examined. — See Hale (1972b:344) for records
from the Union of South Africa.
Pseudoparmelia pustulescens
Figure 14/
Pseudoparmelia pustulescens (Kurokawa) Hale, 1974:191.
Parmelia pustulescens Kurokawa in Hale and Kurokawa,
1964:156 [type collection: Vila Flor, Humbo, Angola,
Degelius (Degelius herbarium, holotype; TNS, US, iso-
types)].
Parmelia imperfecta Kurokawa in Hale and Kurokawa, 1964:
155 [type collection: Bergville, Natal, Union of South
Africa, Almborn 8815 (LD, holotype; US, isotype)].
Description. — Thallus as in P. owariensis (see
above). Apothecia adnate, 1-2 mm in diameter, the
amphithecium pustulate; spores 8, 4-5 X 7-9 p,m.
Chemistry; — Medulla K — , C — , KC — , P — ; atra-
norin, sekikaic acid, fatty acids, and unknown sub-
stances.
Distribution. — Africa and India.
Habitat. — On rocks in open areas up to 2000 m
elevation.
Remarks. — This species is essentially indistin-
guishable from P. owariensis (Asahina) Hale and
a chemical test is needed to separate them. They
appear to be sympatric although collected to-
gether only in the Ivory Coast. By using thin-layer
chromatography I had earlier synonymized P. im-
perfecta (Hale, 1972b).
Specimens Examined. — Ivory Coast: Seguela, Santesson
10691b (UPS, US). Guinea: Zerekore, Santesson 10584. Union
of South Africa: Natal, Almborn 6110 (LD), Hoeg (TRH).
India: Tamil Nadu, Hale 43674.
Pseudoparmelia rahengensis
Figure 15a
Pseudoparmelia rahengensis (Vainio) Hale, 1974:191.
Parmelia rahengensis Vainio, 1921:39 [type collection: Nong
Boa, near Ralieng, Thailand, Hosseus 320 (TUR, lecto-
type)].
Description. — Thallus closely adnate on bark or
rock, yellowish green, 3-6 cm broad; lobes sublin-
ear, 0.5-2 mm wide; upper surface convex, contin-
uous, densely isidiate, the isidia mostly simple, to
O. 2 mm high; lower surface brown to tan, moder-
ately rhizinate, the rhizines tan. Apothecia adnate,
1-3 mm in diameter, the amphithecium sparsely
isidiate; spores 8, 3 X 5 pm.
Chemistry. — Cortex K+ yellowish, medulla K — ,
C— , or C T yellow, KC+ orange, P— or P+ faint;
barbatic acid, obtusatic acid, 4-0-demethyl-barbatic
acid, unidentified substance, rarely with an uniden-
tified pigment.
Distribution. — Thailand.
Habitat. — On tree bark or rocks in open decid-
uous forests at 300-1350 m.
Remarks. — This species is unique in producing
barbatic acid. Is is superficially similar to two other
narrow lobed, usnic acid-containing Asian species,
P. dahlii Hale (lecanoric acid), and P. malaccensis
(Nylander) Hale (protocetraric acid). Chemical
tests should be made to separate them.
Specimens Examined. — Thailand: Kerr L28, (BM, US), Kuro-
kawa 1602, 1815 (TNS, US).
Pseudoparmelia raunkiaeri
Figure 156
Pseudoparmelia raunkiaeri (Vainio) Hale, 1974:191.
Parmelia raunkiaeri Vainio, 1915:19 [type collection: Cane
Bay, St. Croix, Raunkiaer 461 (TUR, Vainio herbarium
number 2752, lectotype; C, FH, isotypes)].
Parmelia scabrosa Vainio, 1896a:33 [type collection: Chateau
Belair, St. Vincent, Elliott 249 (TUR, lectotype; BM, iso-
lectotype); not Parmelia scabrosa Taylor, 1847:162 (= Xan-
thoparmelia scabrosa (Taylor) Hale)].
Parmelia vincentina Zahlbruckner, 1929:222 [type collection:
based on Parmelia scabrosa Vainio].
Description. — Thallus closely adnate on bark,
light buff mineral gray, 5-8 cm broad; lobes sub-
irregular, apically rotund, 2-4 mm wide; upper sur-
face plane, becoming irregularly rugose toward the
center, pustulate-sorediate, the pustules coalescing
in a coarsely sorediate mass in older portions; lower
surface black and sparsely rhizinate except for a
narrow bare, brown zone at the tips. Apothecia un-
known.
Chemistry. — Cortex K+ yellow, medulla K — ,
C — , KC — , or C+ KC+ rose; atranorin and pro-
tocetraric acid with or without gyrophoric acid and
unidentified fatty acids.
NUMBER 31
43
Figure 15. Species of Pseudoparmelia: a, P. rahengensis ( Kurokawa 1602 in US); b, P. raun-
kiaeri ( Evans in US); c, P. rodriguesiana ( Santesson 105 60a in US); d, P. rupicola ( Reitz and
Klein 16113 in US); e, P. rutidota ( Heller 260 in US); f, P. salacinifera (Moore 4406 in US).
(Scale in mm except for P. rahengensis which is x3.)
44
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Distribution. — West Indies and Mexico.
Habitat. — On trees in dry, open or secondary
forest at sea level to 500 m elevation.
Remarks. — The soredia of this species are rather
coarse and originate from small pustulate-isidiate
growths on the upper surface. These burst open
and coalesce so that the major part of the thallus
is sorediate. The relationship to another West In-
dian species, isidiate P. martinicana (Nylander)
Hale, is very close. The chemistry is variable. Speci-
mens from St. Croix (including the lectotype) and
Grand Cayman lack gyrophoric acid. All others con-
tain both protocetraric and gyrophoric acids, as does
P. martinicana.
Specimens Examined. — Mexico: Tamaulipas, Pursell 5197
(US). Cuba: Oriente, Hioram 5545 (US). Grand Cayman:
Imshaug 24480, 24449 (MSC, US), 24508 (MSC). Jamaica:
Imshaug 15000, 15571 (MSC), 1565S, 16044 (MSC, US).
Haiti: Sud, Imshaug 23119, 23140 (MSC). Dominican Repub-
lic: Santiago, Wetmore 3910 (MSC). St. Croix: Britton 77
(NY, US), Evans (US, YU), Raunkiaer 461 (C). Tortola: Fish-
lock 495 (FH, NY). St. Barthelemy: Le Gallo 426, 520, 547,
584 (US), 2626 (MSC). Monserrat: Evans 46 (US, YU) Guade-
loupe: Degelius (Degelius herbarium).
Pseudoparmelia rodriguesiana
JlCURE 15c
Parmelia rodriguesiana Hue, 1899:167 [type collection: Am-
bositra, Madagascar, Rodriguez (PC, lectotypes)].
Description. — Thallus closely adnate on rock,
whitish mineral gray, 5-10 cm in diameter; lobes
sublinear to subirregular, 2-3 mm wide; upper sur-
face plane to rugulose, cracked with age; lower sur-
face black, sparsely rhizinate except for a narrow
naked zone near the tips. Apothecia common, ses-
sile, 2-8 mm in diameter; spores 8, 5-7 X 8-1 3 um.
Chemistry. — Cortex K+ yellow, medulla K — ,
C — , KC+ faint purple violet or KC — ; P — ; atra-
norin and divaricatic acid with associated un-
knowns.
Distribution. — Guinea, Ivory Coast, Angola,
Union of South Africa, and Madagascar.
Habitat. — On open or exposed rocks at 1000-
1700 m elevation.
Remarks. — This rarely collected lichen would
seem to be closely related to P. nairobiensis (Steiner
and Zahlbruckner) Hale, which is normally corti-
colous and somewhat larger and more loosely
adnate. Further field studies will be needed to deter-
mine the exact relationship between these two spe-
cies. Pseudoparmelia rupicola (Lynge) Hale in
South America is also saxicolous and contains divar-
icatic acid, but it has smaller, closely appressed
lobes.
Specimens Examined. — Guinea: N’Zerekore, Santesson
10560a (UPS, US). Ivory Coast: Man, Santesson 10635 bis
(UPS, US). Angola: Huambo, Degelius (Degelius herbarium,
US). Union of South Africa: Natal, Almborn 8594 (US);
Transvaal, Kofler (LD).
Pseudoparmelia rupicola
Figure 15 d
Pseudoparmelia rupicola (Lynge) Hale, 1974:191.
Parmelia rupicola Lynge, 1914:132 [type collection: Porto
Alegre, Rio Grande do Sul, Brazil, Malme 1339 (S, lecto-
type)].
Description. — Thallus closely adnate on rock,
whitish to ivory mineral gray, 4-8 cm broad; lobes
sublinear, contiguous, 1-2 mm wide; upper surface
plane to rugulose, shiny, transversely rimose with
age, becoming lobulate toward the center; lower
surface black except for a dark brown zone at the
tips, moderately rhizinate, the rhizines black. Apo-
thecia common, adnate, 1-1.5 mm in diameter;
spores 8, 6 X 7-8 p.m.
Chemistry. — Cortex K-f yellow, medulla K — ,
C — , KC— or KC+ wine colored, P — ; atranorin
and divaricatic acid with associated unknowns.
Distribution. — Brazil and Uruguay.
Habitat. — On sandstone outcrops in open areas
at about 1000 m elevation.
Remarks. — Pseudoparmelia rupicola closely re-
sembles sorediate P. alabamensis (Hale and Mc-
Cullough) Hale in lobe configuration and adnation
on sandstone, and they may be remotely related.
Strictly saxicolous species of Pseudoparmelia are
rare in New World.
Specimens Examined. — Brazil Catarina, Reitz and Klein
16111, 16113, 16115, 16121 (US). Uruguay: Lavalleja, Osorio
6425 (MVM); Rocha, Hosseus (H).
Pseudoparmelia rutidota
Figure 15e
Pseudoparmelia rutidota (Hooker and Taylor) Hale, 1974:
191.
Parmelia rutidota Hooker and Taylor, 1844:645 [type collec-
tion: Van Diemen’s Land, Australia (FH, lectotype)].
Parmelia caperata var. caperatula Nylander, 1860:377 [type
NUMBER 31
45
collection: Australia (H, Nylander herbarium number
35731, lectotype; FH, isolectotype)].
Parmelia caperata f. ramealis Nylander, 1861:373 [type col-
lection: Andes, Bolivia, Mandon (H, Nylander herbarium
number 35692, lectotype; FH, S, isolectotypes)].
Parmelia jelinekii Krempelhuber, 1868:321 [type collection:
Australia, Jelinek 27 (M, lectotype; W, isolectotype)].
Parmelia ochroleuca Muller Argoviensis, 1882:306 [type col-
lection: Near Illawarra, Australia, Kirton 1 (G, lectotype);
not Parmelia ochroleuca Taylor, 1848:24 (= Sticta)].
Parmelia splendidula Delise ex Nylander, 1885:605 [type
collection: Peru (H, Nylander herbarium number 35733,
lectotype)].
Parmelia caperatula (Nylander) Nylander, 1885:606.
Parmelia subcaperatula Nylander, 1885:606 [type collection:
Derwent River, Tasmania, Brown (H, Nylander herbarium
number 35730, lectotype; BM, isolectotype)].
Parmelia confertula Stirton, 1899:77 [type collection: Bris-
bane, Australia, Bailey (BM, lectotype)].
Description. — Thallus adnate to appressed on
bark, light greenish yellow, 3^8 cm broad; lobes sub-
irregular, apically rotund, contiguous, 2-4 mm
wide; upper surface plane or becoming rugulose
and cracked on older lobes; medulla white but
sometimes with a reddish pigment near the lower
cortex; lower surface black, shiny, sparsely rhizinate,
usually with a bare dark brown naked zone at the
tips. Apothecia common, sessile, the rim crenate,
1-3 mm in diameter; spores 8, 7-10 X 14-20 pm.
Chemistry. — Cortex K— or K+ yellowish, me-
dulla K — , C — , KC— or KC+ rose, P+ red; usnic
acid, rarely atranorin, protocetraric acid with or
without associated unknowns, and with or without
caperatic acid (and protolichesterinic acid?) and
unidentified K+ purple pigments.
Distribution. — United States, Mexico, South
America, and Australia.
Habitat. — On trunks and branches of trees in
semiarid regions at 100-2000 m elevation.
Remarks. — Pseudoparmelia rutidota is a rather
variable species, as one might assume from the long
list of synonyms. It occurs very commonly in certain
semiarid regions, especially Texas in the United
States and New South Wales in Australia. Outside
of this it is relatively rare. The comparable African
populations appear to consist entirely of P. am-
plexa (Stirton) Hale, a smaller, more congested
species with a black velvety lower surface. On the
other hand, some specimens from Mexico and South
America are quite a bit larger than the average.
Obviously environmental modification plays a role
in this variation. Basically P. rutidota could be, or
closely resemble, a now extinct progenitor (or pro-
genitors?) of the sorediate morph P. caperata (L.)
Hale, the pustulate morph P. baltimorensis (Gyel-
nik and Foriss) Hale, and the isidiate morph P.
papillosa (Gyelnik) Hale. None of these fit their
respective roles perfectly but together they form a
coherent species group.
Chemical variation centers around the presence
or absence of fatty acids, in most cases caperatic
acid, and a medullary pigment. There seems to be
no geographic pattern here, but assuredly many
more field studies and collections are needed to
determine this. For the present a rather broad spe-
cies concept seems unavoidable.
Specimens Examined. — United States: Texas, Darrow 4863
(US), Hale 5316, 5504, Heller in Lichenes boreali-americani
197 and Decades of North American Lichens 260 (US),
Hubricht B1891, B1927, Jermy (US), Jones (US), Orchard 4
(US), Slater 3 (US), Whitehouse 2283, 2289, 2290 (US). Mex-
ico: Tamaulipas, Nakanishi 184 (US), Purcell 5564, 5601 (US).
Bolivia: Mandon (BM). Brazil: Pernambuco, Xavier 776,
815, 816, 826. Chile: Valparaiso, Follmann 13013 (US). Uru-
guay: Lorentz 740 (M); Minas, Herter 90669 (H). Argentina:
Lorentz (M). Australia: New South Wales, Degelius A-37,
A-38 (Degelius herbarium, US), Doing (L, US), Du Rietz 683,
693 (UPS, US), Hamilton L1812 (NSW), Watts L1817 (NSW);
South Australia, Rogers 1463, 1697 (US); Western Australia,
Irvine (G); Tasmania, Pearcey (BM); Victoria, Filson 5419
(US); Australian Capital Territory, Weber in Lichenes Ex-
siccati 268 (US).
Pseudoparmelia salacinifera
Figure 15/
Pseudoparmelia salacinifera (Hale) Hale, 1974:191.
Parmelia salacinifera Hale in Hale and Kurokawa, 1964:157
[type collection: Sanford, Seminole County, Florida, Rapp
(US, holotype; FLAS, isotype)].
Description. — Thallus adnate on bark, light ashy
buff, 6-12 cm broad; lobes subirregular, apically
subrotund, 3-5 mm wide; upper surface plane to
rugulose, fissured with age, moderately isidiate, the
isidia simple, to 0.3 mm high; lower surface brown
to tan, moderately rhizinate except for a narrow
naked zone along the margins. Apothecia rare, 2-4
mm in diameter, the amphithecium isidiate; spores
8, 8-9 X 13-16 pm.
Chemistry. — Cortex K-+- yellow, medulla K +
yellow turning red, C — , KC — , P+ pale orange;
atranorin and salazinic acid.
Distribution. — Southeastern United States, Mex-
ico, Central America, West Indies, Colombia, Vene-
zuela, Brazil, and Thailand.
46
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Habitat. — On trees (deciduous trees, palm, con-
ifers) in open or secondary forest from sea level to
1000 m elevation.
Remarks. — The distinguishing features of this
species are the isidia and pale brown lower surface.
The only other comparable salazinic acid-containing
species is saxicolous P. scotophylla Kurokawa from
Australia. It has a smaller thallus and small spores.
Pseudoparmelia cinerascens (Lynge) Hale has a
black lower surface. The only other species that
could be confused with it because of the similar
habitat, range, thallus color, and lobe configuration
is P. amazonica (Nylander) Hale, which has a
black lower surface and contains protocetraric acid
(K— )•
Specimens Examined. — United States: Georgia, Hale 16843;
Florida, Hale 7994, 16714, 17045, 17083, 17735, 21690, 21967,
Moore 4406 (DUKE, US). Rapp (FLAS, US), Standley 13084
(US). See Moore (1968:225) for further records from Florida.
Merico: Chiapas, Hale 20607. Honduras: Comayagua, Standley
6522 (F, US); Cortes, Morton 7896 (US). Cuba: Pinar del Rio,
Imshaug 25215 (MSC, US). Jamaica: Plitt (US). Colombia:
Santander, Nee and Mori 3736 (US). Venezuela: Maracay,
Nash 1946 (US). Brazil: Maranhao, Eiten 4198 (US); Mato
Grosso, Malme (UPS). Thailand: Kurokawa 1931 (TNS, US).
Pseudoparmelia schelpei
Figure 16a
Pseudoparmelia schelpei (Hale) Hale, 1974:191.
Parmelia schelpei Hale, 1972b:344 [type collection: Maxixe,
Sul do Save, Mozambique, Schelpe 4460 (BOL, holotype;
LD, US, isotypes)].
Description. — Thallus closely adnate on bark,
whitish mineral gray, 2-4 cm broad; lobes sublinear,
crowded, 1. 5-2.0 mm wide; upper surface plane,
dull, rugulose with age; lower surface black,
sparsely rhizinate. Apothecia numerous, adnate, 1-2
mm in diameter; spores 8, 4 X 8-10 p,m.
Chemistry. — Cortex K+ yellow, medulla K — ,
C — , K— or KC+ rose, P+ orange; atranorin and
protocetraric acid.
Distribution. — Mozambique.
Habitat. — On coconut palms near sea level.
Remarks. — No other species in Pseudoparmelia
has protocetraric acid and such narrow lobes. Pseu-
doparmelia caribaea (Hale) Hale, a saxicolous
species in the West Indies, is much larger. The
sorediate morph of P. schelpei is presumed to be
P. epileuca (Hale) Hale, which also occurs in Mo-
zambique as well as in Kenya. I suspect both species
will be more frequently collected as lichenologists
visit coastal localities in East Africa.
Specimens Examined. — Mozambique: Mogg 2099 (PRE, US).
Pseudoparmelia schistacea, new combination
Parmelia schistacea Kurokawa and Filson, 1975:44 [type col-
lection: 65.5 km west of Kingoonya, South Australia, Filson
11921 (MEL, holotype) (not seen)].
Description. — Thallus closely adnate, pale oliva-
ceous gray, 2-4 cm broad; lobes sublinear-elongate,
crowded at the center of the thallus, 0.5-1. 5 mm
wide; upper surface plane to convex, shiny, be-
coming tangentially rimose, pustulate; medulla
white; lower surface pale brown, sparsely rhizinate.
Apothecia not seen.
Chemistry. — Cortex K+ yellow, medulla K — ,
C — , KC — , P — ; atranorin, a trace of usnic acid,
caperatic acid, and an unidentified fatty acid.
Distribution. — Australia.
Habitat. — On rocks in open areas.
Remarks. — The authors of this species compare
it with P. arcana (Kurokawa) Hale, both having
closely adnate, almost subcrustose thalli. They have
different morphologies, however, P. arcana being
typically isidiate. A trace of usnic acid is alleged to
occur in P. schistacea. The species is illustrated in
Kurokawa and Filson (1975, pi. 4: fig. 1).
Pseudoparmelia scotophylla, new combination
Figure 165
Parmelia scotophylla Kurokawa in Kurokawa and Filson,
1975:45 [type collection: Ardglen Gap, Liverpool Range,
New South Wales, Australia, Kurokawa 5174 (TNS, holo-
type; MEL, isotype) (not seen)].
Description. — Thallus closely adnate, whitish to
dark mineral gray, 4-12 cm broad; lobes sublinear,
congested, apically subrotund, 1-2 mm wide; upper
surface plane, shiny, becoming densely isidiate, the
isidia cylindrical, simple, the tips blackened; lower
surface dark brown or blackening, sparsely to mod-
erately rhizinate, the rhizines brown or black, sim-
ple. Apothecia (from type description) substipitate,
6 mm in diameter, the amphithecium isidiate;
spores 8, 5 X 7-8 pm.
Chemistry. — Cortex K+ yellow, medulla K +
yellow turning red, C — , KC— , P+ orange; atra-
norin and salazinic acid.
NUMBER 31
MR f.
> . a. ' ’ 'tv-' * . • - /g? * i
Figure 16. — Species of Pseudoparmelia: a, P. schelpei ( Schelpe 4460, isotype in US); b. P.
scotophylla (Weber L-47301 in US); c, P. scrobicularis (Malme 1949, isotype of Parmelia
longiconida Lynge in US); d, P. somaliensis (Jellicoe 40 in US); e, P. soredians (Sampaio 247
in US); f, P. sphaerospora (Hale 21970A). (Scale in mm.)
48
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Distribution. — Australia.
Habitat. — On rocks in dry shrubby sclerophyll
woodlands.
Remarks. — This species is part of the Pseudopar-
melia molybdiza (Nylander) Hale complex, char-
acterized by isidia and salazinic acid. The color of
the lower surface seems to be variable, dark brown
and blackening or remaining rather pale. A closely
related nonisidiate species and possible progenitor
is P. spodochroa (Kurokawa and Filson) Hale,
which usually contains both salazinic and norstictic
acids.
Specimens Examined. — Australia: New South Wales, Du
Rietz 389 (UPS, US).
Psendoparmelia scrobicularis
Figure 16c
Pseudoparmelia scrobicularis (Krempelhuber) Hale, 1974:191.
Parmelia scrobicularis Krempelhuber, 1873:10 [type collection:
Lagoa Santa, Brazil, Warming (M, lectotype)].
Parmelia longiconida Lynge, 1914:130 [type collection: Rio
Apa, Colonia Risso, Paraguay, Malme 1949 (S, lectotype: LD,
US, W, isolectotypes)].
Description. — Thallus closely adnate on bark,
light buff mineral gray, 2-5 cm broad; lobes sub-
linear, contiguous, 1 .0-1.5 mm wide; upper surface
finely foveolate, dull to shiny, lower surface black,
moderately rhizinate. Apothecia common, adnate,
1.5-2. 5 mm in diameter, the disc white pruinose;
spores 8, 6-7 X 10-12 jxra.
Chemistry. — Cortex K+ yellow, medulla K +
yellow, C — , KC — , P+ orange; atranorin, stictic
acid, and constictic acid.
Distribution. — South America.
Habitat. — On trees in open or secondary forest.
Remarks. — The reticulate wrinkles are very
strongly developed but are not visible without low
power magnification. Two possibly related species,
P. carneopruinata (Zahlbruckner) Hale, a pre-
sumptive sorediate morph, and P. crozalsiana
(Bouly de Lesdain) Hale, have broad reticulately
ridged lobes.
Specimens Examined.- — Venezuela: Distrito Federal, Ernst
(G). Brazil: Minas Gerais, Henschen (UPS). Paraguay: Malme
(UPS): Paraguara, Balansa (G). Argentina: Misiones, Montes
10089 pro parte (LD).
Pseudoparmelia somaliensis
Figure 16d
Pseudoparmelia somaliensis (Muller Argoviensis) Hale, 1974:
191.
Parmelia somaliensis Muller Argoviensis, 1885:501 [type col-
lection: Somalia, Hildebrandt (G, lectotype)].
Parmelia scottii Vainio, 1898a:40 [type collection: Ruwenzori,
Africa, Scott-Elliott (TUR, Vainio herbarium number 2693,
lectotype)].
Parmelia ramulicola Dodge, 1959:172 [type collection: Mad-
agascar, Hildebrandt (FH, holotype)].
Description. — Thallus adnate on twigs, whitish
mineral gray, rather soft and appearing inflated,
3-5 cm broad; lobes subirregular, 2-5 mm wide;
upper surface plane to rugulose, dull, becoming
white-pruinose; lower surface dark brown to black
and moderately rhizinate except for a brown naked
zone along the margins. Apothecia common, sub-
stipitate, almost urceolate, 2-5 mm in diameter,
the amphithecium rugose, pruinose; spores 8, 5-7
X 9-13 p,m.
Chemistry. — Cortex K+ yellow, medulla K — ,
C— , KC-f rose, P+ red; atranorin and protoce-
traric acid.
Distribution. — Central Africa and Madagascar.
Habitat. — On twigs and branches of trees in
open areas up to 2300 m elevation.
Remarks. — The most characteristic habitat of this
species is small twigs. The thallus is thick and soft
with large apothecia. There are no close relatives in
the genus for this African endemic.
Specimens Examined. — Zambia: Jellicoe 40 (BM, US). Tan-
zania: Hdeg (TRH).
Pseudoparmelia soredians
Figure 16e
Pseudoparmelia soredians (Nylander) Hale, 1974:191.
Parmelia soredians Nylander, 1872:426 [type collection: For^a,
Reale, Spain, Nylander (H, Nylander herbarium number
34690, lectotype)].
Parmelia conspersa var. polyphylla f. sorediosa Muller Argo-
viensis, 1891:378 [type collection: San Miguel, Azores, God-
man (G, lectotype)].
Parmelia san-miguelii Gyelnik, 1931b:288 [type collection:
based on P. conspersa var. polyphylla t. sorediosa Muller
Argoviensis].
Parmelia halmaiana Gyelnik, 1935:47 [type collection: Ber-
gues, France, Bouly de Lesdain, in Zahlbruckner Lichenes
Rariores Exsiccati 236 (BP, holotype; BM, BPI, MICH,
UPS, isotypes)].
NUMBER 31
49
Description. — Thallus adnate on branches and
trunks of trees, rarely on rocks, yellowish green,
6-15 cm broad; lobes subirregular, apically rotund,
2-5 mm wide; upper surface plane, shiny, becoming
rugose toward the center, sorediate, soralia orbicu-
lar, coalescing, often occurring along ridges and
wrinkles; lower surface black except for a marginal
brown zone, sparsely rhizinate, the rhizines black.
Apothecia rare, adnate; spores not seen.
Chemistry. — Cortex K — , medulla K+ yellow
turning red, C — , KC — , P+ orange; usnic acid and
salazinic acid.
Distribution. — Europe, Central and South Af-
rica, Argentina, Chile, and New Zealand.
Habitat. — On trees and rarely on rocks in open
forests.
Remarks. — This species is similar to P. caperata
(L.) Hale except for the discrete, orbicular soralia
and different chemistry ( P . caperata is P+ red
with protocetraric acid) . It is totally absent from
North America and eastern Asia, where P. caperata
is very common. In England it behaves as a Medi-
terranean species, occurring mostly in the south-
eastern parts of the country where rainfall is low
and there are no severe frosts (James and Rose,
1973).
Specimens Examined. — Europe: Ireland, Mitchell (M); Eng-
land, Holmes (BM); France, Crozals (US), Dahl (WIS), des
Abbayes in Lichenes Armorici Spectabiles IS (LD), Mangillon
668 (US), Santesson 10077 (UPS); Portugal, Persoon (UPS),
Sampaio in Lichenes de Portugal 247 (LD, M, US), Tavares
in Lichenes Lusitaniae selecti exsiccati 140 (H, LD, M, US,
WIS); Italy, Sbarbaro (US), Sbarbaro in Lichenes Selecti Exsic-
cati 24 (H, LD, M, US). Kenya: Central Province, Maas Geester-
anus (L). Union of South Africa: Transvaal, Almborn 6603,
7501, 7505 (LD), Hdeg (TRH); Natal, Almborn 9641, 9763
(LD), Hdeg (TRH); Basutoland, Kofler (LD); Cape Province,
Almborn 1604, 1997, 1998, 2030, 2099, 2100, 2153, 2121, 4108,
4735, 5361, 10662, 10862 (LD), Hdeg (TRH), Kofler (LD),
Maas Geesteranus 6623 (L), Penfield 220 (PRE). Argentina:
Buenos Aires, Santesson 48 (S); Chubut, Santesson 195 (S).
Chile: Santiago, Mahu 1117, 2034 (US); Concepcidn, Barros
3, 193 (H). New Zealand: Scott 323 (BM).
Pseudoparmelia sphaerospora, new combination
Figure 16/
Parmelia sphaerospora Nylander, 1859:254 [type collection:
Madagascar, (H, lectotype; PC, isolectotype)].
Parmelia leucochlora Tuckerman in Nylander, 1860:392.
Parmelia cubensis Nylander, 1885:611.
Parmelia uleana Muller Argoviensis, 1889:506.
Parmelia flavidoglauca Vainio, 1890:65.
Parmelia endoxantha Merrill, 1909:73.
Parmelia bipindensis Dodge, 1959:59.
Parmelia zenkeri Dodge, 1959:74.
[Full citations of the synonyms are given in Hale (1971b:10).]
Description. — Thallus closely adnate on bark,
5-10 cm in diameter, pale greenish or yellowish
mineral gray, often turning chamois in the her-
barium; lobes sublinear-elongate, 2-4 mm wide; up-
per surface plane to convex, more or less maculate,
rugulose to minutely pitted with age; medulla
white to pale yellow; lower surface light tan to pale
olive brown or darkening, rugose, moderately rhiz-
inate, the rhizines simple, tan. Apothecia common,
adnate, 2-3 mm in diameter; spores 8, 6-8 X 7-9
lim.
Chemistry. — Cortex K— or K+ yellowish, me-
dulla yellowish to orange with color reagents; atra-
norin, stictic acid, norstictic acid, gyrophoric acid,
“quintaria” unknowns, and an unidentified yellow
pigment in various combinations. The majority of
specimens contained atranorin and the pigment;
the next largest group (about 20% of the speci-
mens) had atranorin, stictic acid, and the pigment.
The other combinations (atranorin with the “quin-
taria” unknown, with stictic and norstictic acids,
or with gyrophoric acid) occurred in only a few
specimens.
Distribution. — Southeastern United States, Mex-
ico, Central America, West Indies, South America,
and Central Africa.
Habitat. — On shaded tree trunks in mature for-
ests, Taxodium swamps, or orchards from sea level
to 1100 m elevation.
Remarks. — I had previously discussed the occur-
rence of this species in Dominica under the name
Parmelia congruens Acharius (Hale, 1971b: 11).
After making as definitive a summary as possible of
the chemistry, however, I discovered that the lecto-
type of P. congruens contains usnic and hypoproto-
cetraric acids and is better placed in the genus
Xanthoparmelia, leaving P. sphaerospora as the
earliest name for the Pseudoparmelia species. The
medullary color often darkens in old herbarium
specimens. The isidiate morph is P. cyphellata
Lynge.
Specimens Examined. — United States: South Carolina, Cul-
berson 10329 (DUKE, US); Georgia, Egan El-3403, El-6744
(US), Hale 16832; Alabama, Evans 93, 201, 345, 371 (US);
Mississippi, McDaniel L-3 (US), Pursell 3083, 3153, 4181 (US);
50
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Louisiana, Langlois 47 (US); Texas, Jermy (US); Florida,
Calkins 2 (US), Culberson 10893 (DUKE, US), Hale 16728,
16889, 17652, 17738, 21623, 34134, 36829, 36887, Nash 2035
(US), Rapp 19, 2086 (US), Rapp in Cryptogamae Vindobo-
nenses 3654 (US), Rapp in Lichenes Rariores Exsiccati 379
(BM, UPS, US), Standley 73421 (US) (for further records from
Florida see Moore 1968:224). Mexico: Jalisco, Wirth (US);
Chiapas, Hale 20030, 20174. Guatemala: Baja Verapaz, Hale
45998, Kellerman 8045 (US). Panama: Code, Hale 43571;
Panama, Hale 38546. Costa Rica: Tonduz (US). Bahamas:
Brace 6792 (FH, NY). Cuba: Wright 76 (BM, FH, G, NY,
UPS, US); Pinar del Rio, Imshaug 25349, 25376 (MSC). Haiti:
Nord, Leonard 8023 (S, US), 9002 (NY). Dominica: Hale
35636, 35775, 35779. Venezuela: Distrito Federal, Dennis 1548
(BM); Guarico, Goodland 2 (US), Nilson 1500 (UPS). Peru:
San Martin, Allard 2254a, 22255a (US). French Guiana:
Degelius (Degelius herbarium). Brazil: Spruce 130 (BM);
Maranhao, Eiten 3887 (US); Goias, Irwin 32073 (NY, US);
Mato Grosso, Malme in Lichenes Austroamericani 96 (BM,
LD, S, UC, US), 2642 (UC, UPS, US), Ramos 6545 (BM); Sao
Paulo, Schindler 4586 (US). Cameroon: Zenker 4053 (BR).
Zaire: Hoeg (TRH), Louis 6765, 7151, 8482 (BR, US). Guinea:
Santesson 10574 (UPS, US). Angola: Gossweiler 8013a (BM).
Tanzania: Hoeg (TRH). Madagascar: des Abbayes in Lichenes
Madagascarienses 5 (US).
Pseudoparmelia spodochroa, new combination
Figure 17a
Parmelia spodochroa Kurokawa and Filson, 1975:46 [type
collection: Warren Gorge, Flinders Range, South Australia,
R. Filson 11976 (MEL, holotype; TNS, isotype) (not seen)].
Description. — Thallus adnate on rock, whitish to
dark mineral gray, 3-6 cm broad; lobes sublinear,
apically subrotund, contiguous, becoming lobulate
with age, 1.5-2. 5 mm wide; upper surface plane,
transversely rimose with age; lower surface brown
or blackening, moderately rhizinate, the rhizines
brown or black. Apothecia (from Degelius SA-57)
adnate, 2-3 mm in diameter; spores 8, 6 X 8-11
pm.
Chemistry. — Cortex K+ yellow, medulla K +
yellow turning red, C — , KC — , P+ orange; atra-
norin, salazinic acid, and usually norstictic acid.
Distribution. — Australia and Union of South
Africa.
Habitat. — On more or less exposed rocks in dry
woodlands.
Remarks. — This species is represented by only
five collections, which diverge rather widely in sev-
eral characters. The color of the lower surface, for
example, varies from almost entirely black to pale
brown. Chemistry is uniform except for Almborn
4900 and the holotype, which lack norstictic. The
lobe surface is smooth and shiny except for Degelius
SA-57, which has a roughened, almost coarsely
pruinose surface, a trait seen in specimens of other
genera that grow in exposed desert regions. Further
collecting in Africa and Australia will be needed to
decide whether the limits of the species as described
here are too broad.
Specimens Examined. — Australia: Australian Capital Terri-
tory, Weber L-4722G, L-47282 (COLO, US). Union of South
Africa: Cape Province, Almborn 4900 (LD, US), Degelius
SA-57 (Degelius herbarium, US).
Pseudoparmelia subambigua, new species
Figure 176
Description. — Thallus arete adnatus, mollis,
viridi-flavicans, 5-10 cm latus, lobis subirregulari-
bus, brevibus, apice subrotundatis, congestis, 1.0-
3.0 mm latis; superne planus, centrum versus rugo-
sus, dense sorediatus, soraliis orbicularibus, con-
fluentibus; cortex superior 8-10 p,m crassus, epicor-
tex perforatus, stratum gonidiale 20 p,m crassum,
medulla alba, 110-120 p,m crassa, cortex inferior
7-10 jam crassus; subtus niger, sparse rhizinosus,
rhizinis simplicibus, nigris. Apothecia ignota.
Chemistry. — Cortex K — , medulla K — , C + ,
KC+ red, P — ; usnic acid and lecanoric acid.
Holotype. — Chile; Antofagasta, Paposo Que-
brada Guanillo, elevation 880 m, M. Mahu 3405,
12 September 1972 (US; Mahu herbarium, isotype).
Distribution. — Chile.
Habitat. — On shrubs and fence posts in open
areas from sea level to 900 m elevation.
Remarks. — This lichen could easily be misiden-
tified as Parmeliopsis ambigua (Acharius) Nyland-
er. The lower surface is black, however, and the
chemistry distinctive. Larger specimens may also
resemble Pseudoparmelia soredians (Nylander)
Hale, which contains salazinic acid and which may
well be the most closely related species. It is known
only from Chile.
Specimens Examined. — Chile: Coquimbo, Rundel 7230 (US),
Santesson 2519 (S, US); Santiago, Rundel 7323 (US); Valpara-
iso, Santesson 2808 (S, US).
Pseiuloparmelia subamplexa, new species
Figure 17c
Description. — Thallus ut in Pseudoparmelia am-
plexa (Stirton) Hale sed superficie sorediatus, soraliis
NUMBER 31
51
m
■ I TtYTYi TTffTI 1 1 1 1 ill 1 1 1 1 1 1 1
Figure 17. — Species of Pseudoparmelia: a, P. spodochroa ( Weber L-47220 in US); b, P. subam-
bigua ( Santesson 2519 in US); c, P. subamplexa ( Schiitte 34k, isotype in US); d, P. subtiliacea
( Knight 7, lectotype in H); e, P. subtortula (Lye L202A, holotype in BM); f, P. texana (Hale
42248). (Scale in mm.)
52
SMITHSONIAN CONTRIBUTIONS TO BOTANY
orbicularibus, discretis vel pro parte confluen-
tibus; cortex superior 15-18 pin crassus, epicortatus,
epicortice perforato, stratum gonidiale 15-20 pm
crassum, medulla alba, 11 0-1 BO pm alta, cortex in-
ferior 14—20 p m crassus; subtus niger, opacus, modice
rhizinosus, rhizinis simplicibus. Apothecia ignota.
Chemistry. — Cortex K — , C— , KC— or KC +
rose, P+ red; usnic acid and protocetraric acid.
Holotype. — On tree, Chishawasha, Salisbury Dis-
trict, S. Rhodesia, K. Schutte 34k, 30 December 1953
(LD; US, isotype).
Distribution. — Rhodesia.
Habitat. — On trees in arid scrub land.
Remarks. — This is the sorediate morph of P.
amplexa (Stirton) Hale. It has the same peculiar
black velvety lower surface. As with the parent
species, it is restricted to Africa.
Specimens Examined. — Rhodesia, Hoeg (TRH, US), Schutte
42 (LD, US).
Psendoparmelia subtiliacea
Figure 17d
Pseudoparmelia subtiliacea (Nylander) Hale, 1974:191.
Parmelia subtiliacea Nylander, 1885:614 [type collection: New
Zealand, Knight (H, Nylander herbarium number 35176,
lectotype; UPS, isolectotype)].
Parmelia tiliacea var. feracissima Muller Argoviensis, 1886:
256 [type collection: Guntawang, New South Wales, Aus-
tralia, Hamilton 5 (G, lectotype)].
Description. — Thallus adnate on twigs, whitish
mineral gray, 3-5 cm broad; lobes subirregular,
black rimmed, 1.5-3 mm wide; upper surface plane
to rugulose or foveolate; lower surface black, sparse-
ly rhizinate. Apothecia common, substipitate, 1-5
mm in diameter; spores 8, 6-7 X 12-14 pm.
Chemistry. — Cortex K+ yellow, medulla nega-
tive with all reagents; atranorin and caperatic acid.
Distribution. — Australia and New Zealand.
Habitat. — On twigs of subalpine shrubs and in
dry scrub land.
Remarks. — Pseudoparmelia subtiliacea is a very
rare lichen and as a consequence it is poorly under-
stood. The only close relative seems to be another
Australian lichen, P. corrugativa (Filson and Kuro-
kawa) Hale, which has a pigment in the lower half
of the medulla.
Pseudoparmelia subtortula
Figure lie
Pseudoparmelia subtortula (Hale) Hale, 1974:191.
Parmelia subtortula Hale, 1973b:3 [type collection: E. Mengo
District, Uganda, Lye L202A (BM, holotype; US, isotype)].
Description. — Thallus as in P. tortula (Hale)
Hale (see below) except the upper surface isi-
diate, the isidia cylindrical, simple. Apothecia not
seen.
Chemistry.— Cortex K+ yellow, medulla K — ,
C — , KC + rose, P — ; atranorin, norlobaridone, and
neoloxodic acid.
Distribution. — Uganda and Kenya.
Habitat. — On rocks in open areas at about 1000
m elevation.
Remarks. — Pseudoparmelia subtortula is the pre-
sumptive isidiate morph of P. tortula, a species with
identical chemistry, which occurs chiefly in South
Africa. Both are saxicolous African endemics unique
in producing norlobaridone and neoloxodic acid.
Specimens Examined. — See Hale (1973b) for records from
Uganda and Kenya.
Pseudoparmelia texana
Figure 17/
Pseudoparmelia texana (Tuckerman) Hale, 1974:191.
Parmelia texana Tuckerman, 1858:424 [type collection:
Blanco, Texas, Wright (FH, lectotype; M, US, isolecto-
types)].
Parmelia sublaevigata var. texana (Tuckerman) Nylander in
Triana and Planchon, 1867:307.
Parmelia confluescens Nylander, 1875:725 [type collection: St.
Paul (H, Nylander herbarium number 35178, lectotype;
BM, PC, UPS, isolectotypes)].
Parmelia cingalensis Stirton, 1876:159 [type collection: Galle,
Ceylon (BM, lectotype); illegitimate name],
Parmelia leptophylla Muller Argoviensis, 1891:377 [type col-
lection: Baziya, South Africa, Baer 714 (BM, lectotype; G,
isolectotype)].
Parmelia tenuirima f. sorediata Muller Argoviensis, 1894:258
[type collection: Usambara (Tanganyika), Africa, Holst 787
pro parte (G, lectotype)].
Parmelia exoriens Stirton, 1899:76 [type collection: Brisbane,
Australia, Bailey 215 (BM, lectotype; GLAM, isolectotype)].
Parmelia symmiga Hue, 1899:168 [type collection: Coonoor,
Nilgherries Mountains, India, Gray (PC, lectotype)].
Parmelia subconfluescens Magnusson, 1942:5 [type collection:
Puu Waawaa, north of Hualalai, Hawaii, Selling 5669 (S,
holotype)].
NUMBER 31
53
Parmelia mangenotii des Abbayes, 1951:969 [type collection:
Mankono, Cercle of Seguela, Ivory Coast, des Abbayes
(REN, lectotype; US, isolectotype)].
Parmelia pseudorutidota Asahina, 1952:17 [type collection:
Nagachi, Shinshu, Japan, Takahashi (TNS, lectotype)].
Parmelia tanakae Asahina, 1954:371 [type collection: Kuzu-
Mura, Buzen, Prov. Yamato, Japan, Tanaka (TNS, lecto-
type)].
Parmelia albaniensis Dodge, 1959:121 [type collection: forests
of Albany, Cape of Good Hope, Union of South Africa,
Zeyher 3 (FH-Tayl, holotype)].
Description. — Thallus closely adnate on trees
or rocks, ashy white, 6-12 cm broad; lobes sublinear
to subirregular, apically rotund, 3-5 mm wide; up-
per surface plane to rugulose, usually deeply rimose,
sorediate, the soralia laminal, punctiform to capi-
tate, in part initially pustulate with formation of
soredia; lower surface black and moderately rhiz-
inate except for a narrow naked brown zone along
the margins. Apothecia rare, 2-5 mm in diameter;
spores 8, 6-7 X 9-1 1 /un.
Chemistry. — Cortex K+ yellow, medulla K — ,
C— , KC+ pale violet or KC — , P — ; atranorin,
divaricatic acid, and associated unknowns.
Distribution. — Pantemperate outside of Europe
and pantropical at higher elevations.
Habitat. — On trees ( Acacia , Eucalyptus , Junip-
erus, Pinus, Quercus) and more rarely on rocks in
open or secondary forest and in parks and gardens
at 50-2400 m elevation.
Remarks. — This is one of the most widespread
species in the genus and while not usually occur-
ring in as great abundance, for example, as P. cap-
erata (L.) Hale, it is well represented, if not
frequently misidentified, in most large herbaria.
The range of morphological variation is small and
the chemistry uniform and distinctive. It is difficult
to understand why so many lichenologists failed to
recognize it. The nonsorediate progenitor may be
represented by the African P. nairobiensis (Steiner
and Zahlbruckner) Hale and the companion isid-
iate morph by P. concrescens (Vainio) Hale.
Representative Specimens Examined. — United States: Wis-
consin, Thomson 10921 (US, WIS); Tennessee, Hale 37032;
West Virginia, Hale 10308; Illinois, Skorepa 934 (US); Vir-
ginia, Hale 12814 (US); South Carolina, Culberson 7515
(DUKE); Alabama, Hale 34057; Georgia, Hale in Lichenes
Selecti Exsiccati 270 (US); Florida, Hale 36897; Arkansas,
Hale 5567. Mexico: Chihuahua, Matthews 43794 (US); Hi-
dalgo, Chase 7429 (F, US), Weber et al. 33690 (COLO);
Michoacan, Hale 20849; Oaxaca, Cain 27578 (TRT, US),
Hale 20621, 20630; Veracruz, Hale 19407; Chiapas, Hale
20278. Honduras: Morazan, Standley 4105, 11623 (F). Costa
Rica: San Jose, Quiros 1246, 1474 (US). Panama: Chiriqui,
Hale 38914, Scholander (US). Haiti: Ouest, Imshaug 22537
(MSC), Wetmore 2670 (MSC, US). Venezuela: Merida, Hale
42121, 42920. Peru: Abancay, litis 2322 (WIS). Brazil: Minas
Gerais, Eiten 6864, 6870 (US), Mosen 2309 (S, US): Sao Paulo,
Eiten 2978, 8060 (US), Milne 32 (BM), Osorio 4919 (MVM);
Santa Catarina, Reitz and Klein 15114 (US); Rio Grande do
Sul, Lima 90 (US). Uruguay: Canelones, Zorrdn 1929 (US);
Montevideo, Osorio 1849, 6824 (MVM). Chile: Antofagasta,
Rundel 7205 (US). Uganda: Busiro County, Swinscow 2U
24/4 (BM, US). Angola: Bi£, Degelius (Degelius herbarium).
Kenya: Rift Valley Province, Maas Geesteranus 4551 (L).
Rhodesia: Hoeg (TRH). Tanzania: Arusha Prov., Santesson
21338 (UPS); Kilimanjaro Prov., Santesson 20948 (UPS);
Union of South Africa: Natal, Hoeg (TRH); Transvaal,
Almborn 6253 (LD); Cape Province, Almborn 10664 (LD).
Madagascar: des Abbayes in Lichenes Madagascarienses et
Borbonici Selecti Exsiccati 11 (LD, US). India: Tamil Nadu,
Foreau 4092 (Awasthi herbarium). Hale 40167, 43708. Thai-
land: Kurokawa 1948 (TNS). Japan: Province Ohmi, Hale
29473. Indonesia: Java, Groenhart 1711 (L), Kurokawa in
Lichenes Rariores et Critici Exsiccati 39 (TNS, US); Sumatra,
Groenhart 957 (L). Australia: New South Wales, Flockton
724B (US).
Pseudoparmelia tortula
Figure 18a
Pseudoparmelia tortula (Kurokawa) Hale, 1974:191.
Parmelia tortula Kurokawa in Hale and Kurokawa, 1964:157
[type collection: Namaqualand, Cape Province, Union of
South Africa, Almborn 4805 (LD, holotype; TNS, US, iso-
types)].
Description. — Thallus closely adnate on rock,
whitish mineral gray, 5-10 cm in diameter; lobes
subirregular to sublinear, apically subrotund,
crowded toward the center, 1.5-4 mm wide, often
twisted and contorted when free from the substra-
tum; upper surface plane to rugulose or undulate,
continuous, sometimes faintly pruinose; lower sur-
face pale brown, sparsely rhizinate, the rhizines
brown. Apothecia adnate to substipitate, 1-5 mm
in diameter; spores 5-6 X 7-9 /xm.
Chemistry. — Cortex K+ yellow, medulla K — ,
C — , KC— or KC+ faint rose, P — ; atranorin, nor-
lobaridone, and neoloxodic acid.
Distribution. — Union of South Africa.
Habitat. — On rocks in open treeless or sparsely
vegetated areas up to 1000 m elevation.
Remarks. — This rather undistinguished lichen is
still known only from South Africa, where it seems
to be quite common. The presumptive isidiate
54
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Figure 18. — Species of Pseudoparmelia: a, P. tortula ( Almborn 4805, holotype in LD); b,
P. vanderbylii ( Almborn 5110 in US); c, P. venezolana (Hale 45526a, holotype in US); d, P.
violacea (Almborn 1771, holotype in LD); e, P. xanthomelaena (Maas Geesteranus 12145 in US);
/, P. zambiensis (Jellicoe 50, holotype in BM); g, P. zimbabwensis (Hoeg, holotype in TRH).
(Scale in mm.)
NUMBER 31
55
morph, P. subtortula (Hale) Hale, occurs farther
north in Kenya and Uganda.
Specimens Examined. — See Hale and Kurokawa (1964:158)
for records from the Union of South Africa.
Pseudoparmelia vanderbylii
Figure 18&
Pseudoparmelia vanderbylii (Zahlbruckner) Hale, 1974:191.
Parmelia vanderbylii Zahlbruckner, 1932a:252 [type collec-
tion: Union of South Africa, Van Rhynsdorp, Cape Prov-
ince, P. A. van der Byl (W, lectotype)].
Description. — Thallus closely adnate on rocks,
brownish or dark mineral gray, 3-6 cm broad; lobes
subirregular, apically subrotund, contiguous, 1-2
mm wide; upper surface plane, shiny, rimose with
age; lower surface pale brown, sparsely rhizinate,
the rhizines pale brown. Apothecia common, ses-
sile, 1-2 mm in diameter; spores 8, 6 X 8-10 ^m.
Chemistry. — Cortex K+ yellow, medulla K —
or K+ very faint orange, C — , KC — , P — ; atra-
norin and two or more unidentified substances.
Distribution. — Union of South Africa.
Habitat. — On exposed rocks in open areas.
Remarks. — This species falls near P. molybdiza
(Nylander) Hale in general aspect and habitat but
differs in chemistry. Two specimens (the lectotype
and Almborn 4636) have identical chemistry (two
spots in both the hexane and benzene solvent sys-
tems) and the third (Almborn 5110) has distinctly
different but still unidentified higher spots. All
three share the same two lowermost faint spots.
None matched any of the usual KC— depsidones
and depsides with which they were co-chromato-
graphed.
Specimens Examined. — Union of South Africa: Cape Prov-
ince, Almborn 4636, 5110 (LD, US).
Pseudoparmelia venezolana, new species
Figure 18c
Description. — Thallus adnatus, saxicola, cinereo-
albus, 3-7 cm latus, lobis sublinearibus, congestis,
0. 6-1.0 mm latis; superne planus, nitidus, dense
isidiatus, isidiis cylindricis, simplicibus vel ramosis,
0.2-0.4 mm altis; cortex superior 15-18 p,m crassus,
epicorticatus, epicortice distincte perforato, stra-
tum gonidiale ca 10 ^m crassum, medulla alba, 100
p.m crassa, cortex inferior paraplectenchymatus, 8-
10 fxm crassus; subtus pallide brunneolo-albus,
sparse rhizinosus, rhizinis simplicibus, pallidis. Apo-
thecia ignota.
Chemistry. — Cortex K-f yellow, medulla K — ,
C — , KC— or KC+ rose, P+ red; atranorin and
protocetraric acid.
Holotype. — Venezuela: Tachira, Pico Banderas,
Paramo de Tama, elevation 3200 m, M. E. Hale
45526a, 27 March 1975 (US).
Remarks. — This inconspicuous species is easily
overlooked in the high paramo areas where it ap-
pears to be confined. This is the highest elevation
at which the genus is found. There seem to be no
close relatives. It would more likely be confused
with Parmeliopsis, which is characterized by the
presence of thamnolic acid.
Specimens Examined. — Venezuela: Tachira, Hale 45499a,
45523, 45528a.
Pseudoparmelia violacea
Figure 18d
Pseudoparmelia violacea (Kurokawa) Hale, 1974: 191.
Parmelia violacea Kurokawa in Hale and Kurokawa, 1964:
158 [type collection: Blinkwater Ravine, Table Mountain,
Cape Province, Union of South Africa, Almborn 1771 (LD,
holotype)].
Description. — Thallus closely adnate on rock,
greenish mineral gray, 3-6 cm in diameter; lobes
sublinear, 1-3 mm wide; upper surface plane to
distinctly rugulose, continuous; medulla vinaceous
purple; lower surface pale brown, moderately rhiz-
inate, the rhizines pale. Apothecia adnate, 1-1.5
mm in diameter; spores not developed.
Chemistry. — Cortex K+ yellowish, medulla K +
purple, P — ; unidentified anthraquinones.
Distribution. — Union of South Africa.
Habitat.— On rocks in open forest.
Remarks. — -This remarkable species has been re-
collected once since the original description. It is
another typical saxicolous African endemic verging
close to Xanthoparmelia. The pigments are, in fact,
very close to those of X. endomiltoides (Nylander)
Hale, one being identical. No lichen substances are
present in the cortex.
Specimens Examined. — Union of South Africa: Cape Prov-
ince, Degelius SA-70 (Degelius herbarium, US).
56
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Pseudoparmelia xanthomelaena
Figure 18e
Pseudoparmelia xanthomelaena (Muller Argoviensis) Hale,
1974:191.
Parmelia xanthomelaena Muller Argoviensis, 1883a:48 [type
collection: Grampian Mountains, Australia, Sullivan 28
(G, lectotype)].
Description. — Thallus closely appressed on rock,
buff mineral gray but darkening to brownish olive
gray at the center, 5-5 cm broad; lobes sublinear,
contiguous, lobulate, 0.5-1. 5 mm wide; upper sur-
face plane, shiny, transversely rimose with age;
lower surface black, moderately rhizinate. Apo-
thecia common, adnate, up to 1 mm in diameter;
spores 8, 6 X 7-10 ^m.
Chemistry. — Cortex K-f- yellow, medulla K +
yellow, C— , KC — , P+ orange; atranorin, stictic
acid, and constictic acid.
Distribution. — Union of South Africa and Aus-
tralia.
Habitat. — Exposed rocks from sea level to 250
m elevation.
Remarks. — This is one of the smallest species of
Pseudoparmelia. It is probably difficult to collect
since it cannot be removed free of the rock sub-
stratum. There are no obviously related species in
the genus, unless we consider P. ischnoides (Kuro-
kawa) Hale as a possible isidiate morph.
Specimens Examined. — Union of South Africa: Basutoland,
Kofler (LD); Cape Province, Almborn 127, 275, 1136a, 1444,
1546, 1703, 3023, Degelius SA-337 (Degelius herbarium), Maas
Geesteranus 12145 (L).
Pseudoparmelia zambiensis
Figure 18/
Pseudoparmelia zambiensis (Hale) Hale, 1974:191.
Parmelia zambiensis Hale, 1972b:346 [type collection: Nyika
Plateau, Zambia, Jellicoe 59 (BM, holotype; US, isotype)].
Description. — Thallus adnate on bark, greenish
yellow, 4-6 cm broad; lobes subirregular, crowded,
2-3 mm wide, becoming lobulate toward the center;
upper surface plane to rugulose, rimose with age;
lower surface black, sparsely rhizinate, the rhizines
thick, black. Apothecia common, adnate 2. 5-3.0
mm in diameter; spores 8, 6 X 12 jjl m.
Chemistry. — Cortex K — , medulla K — , C — ,
KC— or KC+ faint rose, P— ; usnic acid and di-
varicatic acid with associated unknowns.
Distribution. — Central Africa.
Habitat. — On trees in open areas at 2300 m.
Remarks. — This is another African species that
will surely be collected more often as lichenologists
visit the continent. It appears to be the nonisidiate
progenitor of P. ecaperata (Muller Argoviensis)
Hale, which has a wide range in Africa and Asia.
It could also be considered as the usnic acid chemi-
cal counterpart of P. nairobiensis, which contains
atranorin only in the cortex. A full understanding
of the relations of these species can only be gained
when more specimens are collected.
Pseudoparmelia zimbabwensis
Figure 18g
Pseudoparmelia zimbabwensis (Hale) Hale, 1974:191.
Parmelia zimbabwensis Hale, 1972b:346 [type collection: Zim-
babwe, Rhodesia, Hoeg (TRH, holotype; LD, US, iso-
types)].
Description. — Thallus closely adnate on rock,
fragile, light mineral gray, 2-5 cm broad; lobes sub-
linear, contiguous, black-marginate, 1.0-1. 5 mm
wide; upper surface plane to rugulose, in part
pruinose, pustulate, the pustules thick, apically
pruinose and finally erupting without soredial for-
mation; lower surface black, moderately rhizinate,
the rhizines black. Apothecia not seen.
Chemistry. — Cortex K+ yellow, medulla K — ,
C — , KC — , P+ red; atranorin and protocetraric
acid.
Distribution. — Rhodesia and Union of South
Africa.
Habitat. — On boulders in open areas.
Remarks. — The pustules of this species are simi-
lar to those of P. pusutlescens (Kurokawa) Hale, a
somewhat larger species containing sekikaic acid
(?-)•
Specimens Examined. — Union of South Africa: Transvaal,
Almborn 6108, 6491, 6496 (LD).
Doubtful and Rejected Names
Parmelia appressa Sprengel, 1987:58 [type collec-
tion: On Cinchona , tropical America],
The type collection could not be found. A speci-
men in Acharius’ herbarium labeled “Parmelia
appressa” appears to be identical with Pseudo-
parmelia sphaerospora (Nylander) Hale. How-
NUMBER 31
57
ever, Sprengel mentioned affinity with Parmelia
perforata and P. perlata, both large Parmotrema
species.
Parmelia caperata var. coerulea Bouly de Lesdain,
1954:222 [type collection: Rocamadour, France,
Duval],
The type specimen is probably in PC but I was
not able to locate it there.
Parmelia protorevoluta Gyelnik, 1931b:288.
Parmelio revoluta f. nuda Muller Argoviensis, 1891:378 [type
collection: Santarem, Brazil, Spruce 135 (G, lectotype; BM,
isolectotype)].
The fragmentary type collections contain divari-
catic acid. This is almost certainly a Pseudo-
parmelia, perhaps P. nairobiensis (Steiner and
Zahlbruckner) Hale, a presumed African en-
demic.
Parmelia rutidota var. vestita Zahlbruckner, 1941:
107.
This variety is based on two collections, one
Thomson ZA217 from Southland, New Zealand,
which is Parmelia rudecta Acharius, and the
other Cranwell ZA242 from Coromandel Penin-
sula, New Zealand, which can be identified as
Parmotrema crinitum (Acharius) Choisy. Both
specimens are preserved in W.
Parmelia soredians f. farinosa (Vainio) Gyelnik,
1938a:27.
This is based on Parmelia farinosa Vainio (1890:
62) ( Lichenes Brasilienses Exsiccati 551 in
TUR). It is a species of Xanthoparmelia (Vainio)
Hale.
Parmelia soredians f. lobulata Gyelnik, 1934:306
[type collection: La Calera Sud, Argentina, H os-
sens 22 (BP, holotype)].
This is a species of Xanthoparmelia (Vainio)
Hale.
Parmelia soredians f. muscicola Gyelnik, 1934:306
[type collection: Capilla del Monte, Argentina,
Hosseus 23 (BP, holotype)].
This is also a Xanthoparmelia.
Pseudoparmelia aradensis Gyelnik, 1933:6.
Culberson and Culberson (1969:515) place this
species in synonymy under Cetrelia olivetorum
(Nylander) W. Culberson and C. Culberson.
Pseudoparmelia euplecta (Stirton) Hale, 1974:190.
I have tentatively placed this species as a synonym
of Pseudoparmelia caperata in the “Taxonomic
Treatment.”
Pseudoparmelia leucopis (Krempelhuber) Hale,
1974:190.
Parmelia leucopis Krempelhuber, 1878:461 [type collection:
Lorentz and Hieronymus, Argentina (M. lectotype)].
After reexamining the type collection, I found
some cilia in the lobe axils. These would exclude
the species from Pseudoparmelia, and I hesitate
to reassign it to another genus until better mate-
rial is available.
Pseudoparmelia meiosperma (Hue) Hale, 1974:190.
Parmelia internexa f. meiosperma Hue, 1899:185 [type col-
lection: Mafate, Reunion, Rodriguez (PC, lectotype)].
The type-specimen has rather broad lobes with
dense isidia. The rhizines are dense and more or
less branched. The chemical constituents include
atranorin, evernic acid, and traces of lecanoric
acid, a combination that is known in several spe-
cies of Hypotrachyna (Hale, 1975a) but in no
Pseudoparmeliae. It does not fit well in Pseudo-
parmelia and until additional material is avail-
able it is impossible to settle its generic position.
Pseudoparmelia pseudofallax Gyelnik, 1933:6.
Culberson and Culberson (1968:459) consider
this to be a species of Cetrelia but not having
examined the type collection they could not de-
cide which one.
Pseudoparmelia pseudofallax var. cretaceoides Gyel-
nik, 1933:7.
This taxon also belongs in the Cetrelia cetrari-
oides complex.
Pseudoparmelia pseudosorediosa (Gyelnik) Hale,
1974:191.
This has been reduced to synonymy under Pseu-
doparmelia caperata in the “Taxonomic Treat-
ment” above.
Literature Cited
des Abbayes, H.
1951. Lichens recoltes en Guinee frangaise et en Cote
d’Ivoire. Bulletin d’Institut frangais d’Afrique Noire,
1 3(4) :965— 977.
1958. Lichens recoltes en Guinee franchise et en Cote
d’Ivoire. Bulletin de I’Institut frangais d’Afrique
Noire, 20A(l):l-27.
Acharius, E.
1803. Methodus qua omnes detectos Lichenes. 394 pages.
Stockholm.
1814. Synopsis methodica lichenum. 392 pages. Lund.
Asahina, Y.
1952. Lichenes Japoniae novae vel minus cognitae (9).
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1953. Lichenes Japoniae novae vel minus cognitae (11).
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1954. Lichenologische Notizen (112-113). Journal of Jap-
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Bouly de Lesdain, M.
1923. Notes lichenologiques, XX. Bulletin de la Societe
Botanique de France, 23:277-283.
1954. Notes lichenologiques, No. XXXIX. Bulletin de la
Societe Botanique de France, 101:222-226.
Crombie, J. M.
1876a. New Lichens from the Cape of Good Hope. Journal
of Botany British and Foreign, 14:18-21.
1876b. On the Lichens Collected by Prof. R. O. Cunning-
ham. Journal of the Linnean Society of London,
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NUMBER 31
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SMITHSONIAN CONTRIBUTIONS TO BOTANY
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Index
(Synonyms in italics)
Cetrelia olivetorum, 57
Lichen caperatus, 20
Parmelia adspersa, 14
alabamensis, 14
albaniensis, 53
amazonica, 16
amplexa, 16
angolensis, 38
annexa, 17
antarctica, 32
appressa, 56
aptata, 17
arcana, 17
asmarana, 17
atrichoides, 38
austroafricana, 25
baltimorensis, 18
basutoensis, 18
benguellensis, 16
bipindensis, 49
brachyphylla, 38
caffrorum, 25
capensis, 25
caperata, 20
caperata var. caperatula, 44
caperata var. coerulea, 57
caperata b. cyliphora, 20
caperata var. exornata, 31
caperata var. glaucopis, 16
caperata var. isidiophora, 41
caperata £. laevissima, 20
caperata f. ramealis, 45
caperata var. subglauca, 20
caperatula, 45
caribaea, 21
carneopruinata, 22
caroliniana, 22
chapadensis, 24
cinerascens, 24
cingalensis, 52
citrinescens, 31
concrescens, 24
condyloides, 25
confertula, 45
confluescens, 52
congruens, 49
conspersa var. polyphylla f. sorediosa,
48
corrugativa, 25
crozalsiana, 27
cryptochlorophaea, 27
cubensis, 49
cylisphora, 20
cyphellata, 28
djalonensis, 29
ecaperata, 28
ecoronata, 34
endoxantha, 49
epileuca, 29
er up tens, 29
euplecta, 20, 21
exoriens, 52
ferax, 31
filipina, 14
flavicans, 20
flavidoglauca, 49
ganguellensis , 38
geesterani, 31
gerlachei, 32
glaucopis, 16
gracilenta, 34
gracilescens var. angolensis, 38
gracilis, 34
halmaiana, 48
hansfordii, 39
herreana, 20
hypomilta, 32
imperfecta, 42
inactiva, 14
inhaminensis , 32
inornata, 34
internexa f. meiosperma, 57
intertexta, 34
ischnoides, 35
isidiophora, 22
jelinekii, 45
labrosa, 35
laeteflavens, 29
lecanoracea, 35
leptophylla, 52
leucochlora, 49
leucopis, 57
leucoxantha, 37
longiconida, 48
malaccensis, 37
malaccensis var. laeteflavens, 29
mangenotii, 53
martinicana, 37
molybdiza, 38
nairobiensis, 38
negativa, 20
nipponica, 17
obversa, 31
ochroleuca, 45
ochroleuca f. sorediosa, 20, 21
owariensis, 39
pachydactyla, 41
papillosa, 41
perfissa, 38
perlata var. flavicans, 20
perlata var. subrevoluta, 22
prolata, 41
protorevoluta, 57
pseudorutidota, 53
pseudosorediosa, 20
pustulescens, 42
rahengensis, 42
ramulicola, 48
raunkiaeri, 42
regnellii, 32
regnellii f. arida, 32
relicina var. ecoronata, 34
revoluta f. nuda, 57
rodriguesiana, 44
rupicola, 44
rutidota, 44
rutidota f. filizans, 31
rutidota var. vestita, 57
salacinifera , 45
san-miguelii, 48
sbarbaronis, 22
scabrella, 18
scabrosa, 42
schelpei, 46
schist ace a, 46
scotophylla, 46
scottii, 48
scrobicularis, 48
somaliensis, 48
soredians, 48
soredians f. farinosa, 57
soredians f. lobulata, 57
soredians t. muscicola, 57
sphaerospora, 49
splendidula, 45
spodochroa, 50
steineri, 41
61
62
SMITHSONIAN CONTRIBUTIONS TO BOTANY
subcaperatula, 45
subconfluescens, 52
subconspersa var. benguellensis,
subglauca, 20
sublaevigata var. texana, 52
subrupta, 34
subtiliacea, 52
subtortula, 52
symmiga, 52
tanakae, 53
tenuirima var. labrosa, 35
tenuirima f. sorediata, 52
texana, 52
tiliacea var. feracissima, 52
tortula, 53
uleana, 49
vanderbylii, 55
vincentiana, 42
violacea, 55
wainioana, 22
xanthomelaena, 56
zambiensis, 56
zenkeri, 49
zimbabwensis, 56
Parmotrema crinitum, 57
Pseudoparmelia adspersa, 14
alabamensis, 14
amazonica, 16
amplexa, 16
annexa, 17
aptata, 17
aradensis, 57
arcana, 17
baltimorensis, 18
basutoensis, 18
benguellensis , 16
callichroa, 20
caperata, 20
caribaea, 21
pachydactyla, 41
carneopruinata, 22
papillosa, 41
caroliniana, 22
prolata, 41
chapadensis, 24
pseudofallax, 57
cinerascens, 24
pseudofallax var. cretaceoides, 57
concomitans, 24
pseudosorediosa, 20, 57
concrescens, 24
pustulescens, 42
condyloides, 25
rahengensis, 42
conlabrosa, 25
raunkiaeri, 42
corrugativa, 25
rodriguesiana, 44
crozalsiana, 27
rupicola, 44
cryptochlorophaea, 27
rutidota, 44
cyphellata, 28
salacinifera, 45
dahlii, 28
schelpei, 46
ecaperata, 28
schistacea, 46
epileuca, 29
scotophylla, 46
eruptens, 29
scrobicularis, 48
euplecta, 20, 57
somaliensis, 48
exornata, 31
soredians, 48
ferax, 31
sphaerospora, 49
geesterani, 31
spodochroa, 50
gerlachei, 32
subambigua, 50
hypomilta, 32
subamplexa, 50
inhaminensis, 32
subtiliacea, 52
inornata, 34
subtortula, 52
intertexta, 34
texana, 50
ischnoides, 35
tortula, 53
labrosa, 35
vanderbylii, 55
lecanoracea, 35
venezolana, 55
leucopis, 57
violacea, 55
leucoxantha, 37
xanthomelaena, 56
malaccensis, 37
zambiensis, 56
martinicana, 37
zimbabwensis, 56
meiosperma, 57
Pyxine azorea, 22
molybdiza, 38
Xanthoparmelia gerlachei, 32
nairobiensis, 38
molliuscula, 41
neoquintaria, 39
papillosa, 41
owariensis, 39
scabrosa, 42
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